Oceanography and Marine Biology Repaired

50931_C001.
fm Page 16 Thursday, May 10, 2007 8:38 PM
WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS
2002). Because the VSF is smoother for spheroids (Figure 4B,E, see also Colour Figure 4 in the
insert following p. 344), the anomalous diffraction peaks inherent in spheres determine the pattern
of the biases (Figure 4C,F). Internal transmission and refraction cause the number of peaks in the
VSF for spheres to increase with particle size; however, the magnitudes are dampened and so is
the general trend in the bias.
For both large non-spherical organic-like and inorganic-like particles, forward scattering is
stronger compared with that of equal-volume spheres (Figure 4). In the backward and side-scattering
directions, however, there are differences in the biases in the VSF between the two types of particles;
for organic-like particles, the largest biases are in the backward direction and are associated with
small particles (in particular, particles on the order of the wavelength of light, e.g., D ≈ 0.5 µm;
Figure 4C). For inorganic-like particles the largest differences are in the side-scattering direction
and are associated with large particles (Figure 4F).
Attenuation, absorption and scattering: efficiency factors and biases

Efficiency factors for attenuation, Qc, as a function of particle size, show a similar trend of variation
for spheres and spheroids (Figures 5 and 6), approaching an asymptotic value of two when the GO
limit is reached (Figure 6A,B). The size, D, however, at which Qc reaches its maximal value
increases with increased departure from a spherical shape (Figure 6A,B).
In general, a sphere will overestimate the attenuation (γc < 1) of an equal-volume spheroid (up
to 50% for the most extreme shapes) but will underestimate the attenuation (γc > 1) of an equal-
volume spheroid for particles larger than the wavelength (Figures 7A,C and 8A,B). Scattering
dominates attenuation; the efficiency factors and biases for scattering are very similar to those of
attenuation (Figures 6E,F, 8E,F, 9A,C and 10A,C).
The trend in the change of the efficiency factors for absorption, Qa, as a function of particle
size is similar for spheres and spheroids, approaching an asymptotic value of one at the GO limit
(Figures 5B,D and 6C,D). The absorption efficiency factor of spheroids, however, is always lower
than or equal to that of an equal-volume sphere, regardless of particle size and aspect ratio
(Figure 6C,D). Absorption efficiency factors of inorganic-like particles are low (Figure 6D) and the
biases in absorption between spheres and spheroids are small (Figure 8D). Biases in absorption are
also small for small organic-like particles (γa ≈ 1; Figure 8C), but increase with increasing particle
size and deviation from sphericity. For large organic-like particles, absorption by a spheroid is
always larger than that of a sphere of the same volume (Figures 7B and 8C). That is because the
absorbing material in a randomly oriented spheroid is less packaged compared with that in a sphere,
exposing more absorbing material to the incident light. However, Qa is smaller for a randomly
oriented spheroid (Figures 5B,D and 6C,D), as it is derived from Ca by dividing by the average
cross-sectional area, which is always smaller for spheres.
The backscattering bias can be very large (by a factor of 16, Figure 10B), especially in the
RGD region and for particles much larger than the wavelength (Figures 8G,H and 10B,D). For the
largest particles, the backscattering does not reach the asymptotic value of the other IOPs, at least
in the range of sizes examined here. By applying an unrealistically large absorption value, however,
the backscattering bias does approach the same asymptotic value as total scattering (by using the
T-matrix method, Herring 2002).
The volume-normalised cross sections for attenuation and scattering, αc and αb, respectively,
illustrate that the size contributing most to attenuation and scattering (per unit volume or per unit mass)
is larger for spheroids than for equal-volume spheres (Figure 11A,B,E,F; consistent with the findings
by Jonasz (1987a) that there is as much as a 300% difference between spheres and spheroids in the
volume-normalised attenuation cross section). In general, the magnitude of the volume-normalised
cross sections for attenuation and scattering decreases with departure from sphericity, suggesting
16
50931_C001.fm Page 17 Thursday, May 10, 2007 8:38 PM
INHERENT OPTICAL PROPERTIES OF NON-SPHERICAL MARINE-LIKE PARTICLES
Phase shift parameter ρ Phase shift parameter ρ

0.2 0.5 1 2 5 10 20 50 100 Qc 0.2 0.5 1 2 5 10 20 50 100 Qa
1
A 2.5 B
20
10 0.8
2
5
Aspect ratio ts
1.5 0.6
2
1 1 0.4
0.5
0.5 0.2
0.2
m = 1.05 + i0.01 m = 1.05 + i0.01
0.1
0.5 1 2 5 10 20 50 100 200 Qc 0.5 1 2 5 10 20 50 100 200 Qa

3.5
C D 0.3
20 3
10 0.25
2.5
Aspect ratio ts
5 0.2
2
2 0.15
1.5
1
1 0.1
0.5
0.5 0.05
0.2
m = 1.17 + i0.0001 m = 1.17 + i0.0001
0.1
0.2 0.5 1 2 5 10 20 50 100 200 0.2 0.5 1 2 5 10 20 50 100 200
Particle size D (µm) Particle size D (µm)
Figure 5 Efficiency factors for attenuation, Qc (A, C), and absorption, Qa (B, D), for spheroids as a function
of size, D [µm] (primary x-axis, bottom), with corresponding phase shift parameter, ρ (secondary x-axis, top),
and aspect ratio, s/t. Results were derived using the T-matrix method for small particles (area within the white
line to the left of each plot), and the Paramonov (1994b) method for intermediate and larger particles (rest of
the plot), for two different types of particles: phytoplankton-like particles with refractive index m = 1.05 +
i0.01 (A, B) and inorganic-like particles with m = 1.17 + i0.0001 (C, D).
that spheres interact (resonate) better, per unit volume, with the impinging radiation compared with
other shapes. Similar to attenuation and scattering, model results for the volume-normalised cross
section for backscattering, α bb , show that the size contributing most to backscattering is larger for
spheroids than for equal-volume spheres (Figure 11G,H). Unfortunately, the lack of solutions for
intermediate regions limits the ability here to discuss α bb any further.
The volume-normalised absorption cross section, αa, is higher for all strongly absorbing sphe-
roids with low indices of refraction, consistent with the idea that in a randomly oriented mono-
dispersion of homogeneous spheroids more material can interact with the incident light than in a
monodispersion of spheres of the same volume, which is better ‘packaged’ or ‘self-shaded’
(Figure 11C,D). This is not the case for weakly absorbing particles that are smaller than a few
microns — probably due to scattering within the particle that increases the average path length of
a light ray — hence increasing the probability of absorption. This effect is slightly larger for spheres
given that they are more effective scatterers (Figure 11D).
17

0.2 0.5 1 2 5 10 20 50 0.5 1 2 5 10 20 50 200
3 4
A B
2.5
3
2
Qc
Qc
1.5 2
1
1
0.5
m = 1.05 + i0.01 m = 1.17 + i0.0001
0 0
1.2 0.4
C D m = 1.17 + i0.0001
1
0.3 Aspect ratio st
0.8
10
Qa
Qa
0.6 0.2
2
0.4 1
0.1 0.5
0.2
m = 1.05 + i0.01 0.1
0 0
2.5 4
E F
2
3
1.5
Qb
Qb
2
1
1
0.5
m = 1.05 + i0.01 m = 1.17 + i0.0001
0 0
0.004 0.1
G m = 1.05 + i0.01 H
0.08
0.003
0.06
Q bb
Q bb
0.002
0.04
0.001
0.02
m = 1.17 + i0.0001
0.000 0
0.2 0.5 1 2 5 10 20 50 200 0.2 0.5 1 2 5 10 20 50 200
Figure 6 Efficiency factors for attenuation, Qc (A, B), absorption, Qa (C, D), scattering, Qb (E, F), and back-
scattering, Qbb (G, H), for spheroids as a function of size, D [µm] (primary x-axis, bottom), with corresponding
phase shift parameter, ρ (secondary x-axis, top), and aspect ratio, s/t. Results were derived using the T-matrix
method for small particle sizes and the Paramonov (1994b) method for intermediate sizes, while the ray tracing
method was used to obtain Qbb for large sizes. Results are presented for two different types of particles: a
phytoplankton-like particle with refractive index m = 1.05 + i0.01 (A, C, E, G) and an inorganic-like particle
with m = 1.17 + i0.0001 (B, D, F, H). The lines represent aspect ratios (legend is shown in D): oblate spheroids
with s/t = 0.5 (grey solid line) and s/t = 0.1 (grey dashed line), prolate spheroids with s/t = 2 (dark solid line)
and s/t = 10 (dark dashed line), and spheres with s/t = 1 (solid line with dots).
18

0.2 0.5 1 2 5 10 20 50 100 γc 0.2 0.5 1 2 5 10 20 50 100 γa
3.5
A B 2.75
20
3 2.5
10
2.25
Aspect ratio st
5 2.5
2
2 2
1 1.75
0.5 1.5 1.5
0.2 1 1.25
m = 1.05 + i0.01 m = 1.05 + i0.01
0.1 1
0.5 1 2 5 10 20 50 100 200 γc 0.5 1 2 5 10 20 50 100 200 γa

5
C D 2
20 4.5
4
10 1.8
3.5
Aspect ratio st
5
3 1.6
2 2.5
1.4
1 2
0.5 1.5 1.2
1
0.2 1
m = 1.17 + i0.0001 0.5 m = 1.17 + i0.0001
0.1
0.2 0.5 1 2 5 10 20 50 100 200 0.2 0.5 1 2 5 10 20 50 100 200
Figure 7 Bias in attenuation, γc (A, C), and absorption, γa (B, D). Results were derived as in Figure 5. Thick
grey lines indicate where γ = 1.
The smallest particles (in the Rayleigh limit) have the highest backscattering ratios, b̃ ≈ 0.5
(Figure 12A,C). For particles in the RGD region, the backscattering ratio of non-spherical particles
is largely underestimated by spheres (γ b > 1; Figure 12B,D). The backscattering ratio is lowest in
the transition from the RGD to the VDH regions reaching a constant value for spheres in the mid-
VDH region; b̃ ≈ 0.0005 for organic-like particles and b̃ ≈ 0.0042 for inorganic-like particles. For
large inorganic-like particles, a spherical particle overestimates the backscattering ratio of non-
spherical particles (Figure 12D), though generalisations do not seem possible. The available results
for spheroids, however, are not sufficient to predict how the backscattering ratio of non-spheres
will behave in the intermediate region.
Optical properties of polydispersions

Obtaining the IOPs of polydispersions of particles
When modelling natural waters, it is impractical to account for the contribution of each individual
particle because bulk IOPs are the sums of the IOPs of an assembly of particles varying in size,
composition and shape. In order to model the optical properties of natural populations, assumptions
regarding their size distribution as well as their optical properties need to be made. An advantage
19

0.2 0.5 1 2 5 10 20 50 0.5 1 2 5 10 20 50 200
3 4
A B
2.5
3
2
2
γc
γc
1.5
1
1
m = 1.05 + i0.01 m = 1.17 + i0.0001
0.5 0
2.6 1.8
C m = 1.05 + i0.01 D 10
2.2 1.6 2
Aspect ratio st 1
1.8 1.4 0.5
γa
γa
0.1
1.2
1.4
1
1 m = 1.17 + i0.0001
0.8
4 4
E F
3 3
2 2
γb
γb
1 1
m = 1.05 + i0.01 m = 1.17 + i0.0001
0 0
7 2.5
G H
6 m = 1.05 + i0.01
2
5
4 1.5
γbb
γbb
3 1
2
0.5
1 m = 1.17 + i0.0001
0 0
0.2 0.5 1 2 5 10 20 50 200 0.2 0.5 1 2 5 10 20 50 200
Figure 8 Biases for attenuation, γc (A, B), absorption, γa (C, D), scattering, γb (E, F), and backscattering, γbb
(G, H), for spheroids as a function of size, D [µm] (primary x-axis, bottom), with corresponding phase shift
parameter, ρ (secondary x-axis, top). Each line represents a different aspect ratio, s/t (legend is shown in
panel D). Results were derived as in Figure 6 for two different types of particles: a phytoplankton-like particle
with refractive index m = 1.05 + i0.01 (A, C, E, G) and an inorganic-like particle with m = 1.17 + i0.0001
(B, D, F, H).
20

0.2 0.5 1 2 5 10 20 50 100 Qb 0.2 0.5 1 2 5 10 20 50 100 Qbb
2 0.006
A B
20
0.005
10 1.5
5 0.004
Aspect ratio s
t
2 1 0.003
1
0.002
0.5 0.5
0.2 0.001
m = 1.05 + i0.01 m = 1.05 + i0.01
0.1
0.5 1 2 5 10 20 50 100 200 Qb 0.5 1 2 5 10 20 50 100 200 Qbb

3.5 0.09
C D
20 3 0.08
10 0.07
2.5
5
Aspect ratio s
t
0.06
2
2 0.05
1.5 0.04
1
1 0.03
0.5
0.02
0.2 0.5
m = 1.17 + i0.0001 m = 1.17 + i0.0001 0.01
0.1
0.2 0.5 1 2 5 10 20 50 100 200 0.2 0.5 1 2 5 10 20 50 100 200
Figure 9 Efficiency factors for scattering, Qb (A, C), and backscattering, Qbb (B, D), for spheroids as a function
of size, D [µm] (primary x-axis, bottom), with corresponding phase shift parameter, ρ (secondary x-axis, top),
and aspect ratio, s/t. Results were derived as described in Figure 5. Only the T-matrix and ray tracing methods
were used for Qbb, however, as no other approximation is currently available for intermediate sizes.
of modelling polydispersions is that the results provide more realistic values of IOPs that better
mimic natural populations by eliminating the extreme characteristics of monodispersions that are
averaged out and smoothed in the IOPs for polydispersions.
Particles are distributed according to a particulate size distribution (PSD) that describes how
their number concentration varies with size. Most often the PSD, f (D) [# m–3 µm–1], is given in its
differential form in number of particles per unit volume per unit bin length, such that the number
concentration, N, between two size classes, D1 and D2, is computed as:
∫ f ( D ) dD [ # m
D2
−3
N  ∆D = D2 − D1  = ]. (25)
D1
For a population of particles with varied composition, the IOPs are computed from the PSD as:
∫ ( ) ( )
Dmax
c, a, b, bb = Cc,a,b,bb D f D dD [m −1 ], (26)
Dmin
21

0.2 0.5 1 2 5 10 20 50 100 γb 0.2 0.5 1 2 5 10 20 50 100 γbb
16
A 4.5 B
20 14
4
10 12
3.5
Aspect ratio st
5 10
3
2 2.5 8
1 2 6
0.5 1.5
4
1
0.2 2
m = 1.05 + i0.01 0.5 m = 1.05 + i0.01
0.1
0.5 1 2 5 10 20 50 100 200 γb 0.5 1 2 5 10 20 50 100 200 γbb

5
C D
20 4.5 3.5
4
10 3
3.5
Aspect ratio st
5
3 2.5
2 2.5 2
1 2
1.5
0.5 1.5
1 1
0.2
m = 1.17 + i0.0001 0.5 m = 1.17 + i0.0001 0.5
0.1
0.2 0.5 1 2 5 10 20 50 100 200 0.2 0.5 1 2 5 10 20 50 100 200
Figure 10 Bias in scattering, γb (A, C), and backscattering, γbb (B, D). Results were derived as in Figure 9.
Thick grey lines indicate where γ = 1.
For the analysis conducted here two common PSDs previously used in other studies of IOPs
are employed:
1. The power-law PSD (also referred to as the Junge-like PSD) is given by:
 0, if D < Dmin or D > Dmax ;


( )
f D =
 D 
−ξ [ # m −3µm −1 ], (27)
n0  D0  , if Dmin ≤ D ≤ Dmax
where D0 [µm] is a reference diameter and n0 [ # m −3 µm −1 ] is the differential number

concentration at the reference diameter. ξ is the slope of the differential PSD (values in
aquatic environments are within the range 2.5 < ξ < 5.0; Morel 1973, Jonasz 1983,
Stramski & Kiefer 1991). When ξ = 4.0 the PSD is known as the Junge PSD for which
an equal volume of particulate material is distributed in logarithmically increasing size
bins. The smaller the value of the power-law slope, ξ, the smaller is the relative contri-
bution of small particles to the PSD. In the analysis presented here, ξ is allowed to vary
22

0.2 0.5 1 2 5 10 20 50 0.5 1 2 5 10 20 50 200
1 3.5
A m = 1.05 + i0.01 B m = 1.17 + i0.0001
3
0.8
2.5
0.6 2
αc
αc
0.4 1.5
1
0.2
0.5
0 0
0.3 0.0040
C m = 1.05 + i0.01 D m = 1.17 + i0.0001
0.25
0.0035
0.2
αa
αa
0.15 0.0030
0.1
0.0025
0.05
0 0.0020
0.7 3.5
E m = 1.05 + i0.01 F m = 1.17 + i0.0001
0.6 3
0.5 2.5
0.4 2
αb
αb
0.3 1.5
0.2 1
0.1 0.5
0 0
0.012 0.12
G m = 1.05 + i0.01 H m = 1.17 + i0.0001
0.01 0.1
Aspect ratio st
0.008 0.08
10
αbb
αbb
0.006 0.06
2
0.004 1 0.04
0.5
0.002 0.02
0.1
0 0
0.2 0.5 1 2 5 10 20 50 200 0.2 0.5 1 2 5 10 20 50 200
Figure 11 Volume-normalised cross sections for attenuation, αc (A, B), absorption, αa (C, D), scattering, αb
(E, F), and backscattering, αbb (G, H), for spheroids as a function of size, D [µm] (primary x-axis, bottom),
with corresponding phase shift parameter, ρ (secondary x-axis, top). Each line represents a different aspect
ratio, s/t (legend is shown in panel G). Results were derived as in Figure 6 for two different types of particles:
a phytoplankton-like particle with refractive index m = 1.05 + i0.01 (A, C, E, G) and an inorganic-like particle
with m = 1.17 + i0.0001 (B, D, F, H).
23

0.2 0.5 1 2 5 10 20 50 0.2 0.5 1 2 5 10 20 50
100 8
A 10 B
10−1 Aspect ratio st 2 6 m = 1.05 + i0.01
1
0.5
10−2 4
b̃
γb̃
0.1
10−3 m = 1.05 + i0.01 2
10−4 0
0.5 1 2 5 10 20 50 200 0.5 1 2 5 10 20 50 200

100 5
C D
4 m = 1.17 + i0.0001
10−1 3
b̃
γb̃ 2
10−2 1
m = 1.17 + i0.0001
0
0.2 0.5 1 2 5 10 20 50 200 0.2 0.5 1 2 5 10 20 50 200
Figure 12 Backscattering ratios, b = bb (A, C), and biases in the backscattering ratio, γ b b (B, D), as a function
b
of particle size, D [µm] (primary x-axis, bottom), with corresponding phase shift parameter, ρ (secondary
x-axis, top). Results are derived as in Figure 9 for two different types of particles: a phytoplankton-like particle
with m = 1.05 + i0.01 (A, B) and an inorganic-like particle with m = 1.17 + i0.0001 (C, D). Each line represents
a different aspect ratio, s/t (legend is shown in panel A).
(representing the variations in the natural environment) to examine how changes in the
relative concentration of small to large particles affects biases between spherical and
non-spherical populations of particles.
2. A more elaborate PSD based on generalised gamma functions was introduced by Risoviç
(1993):
 0, if D < Dmin or D > Dmax ;

 µS µL
f ( D) =   D  νS D νL
[# m −3µm −1 ],
nS   exp(− τ S D ) + nL   exp(− τ L D ), if Dmin ≤ D ≤ Dmax
  D0   D0 
(28)
where nS and nL are the number concentrations of small and large particles [# m–3 µm–1],
respectively, and D0 [µm] is the reference diameter. The other parameters, µS,L , τS,L and
υS,L, help to generalise the gamma functions that express the distributions of the small
and large particles, respectively, and are site-specific with values provided by Risoviç
(1993) (parametric values of a ‘typical’ water body are µS = 2, τS = 52 µm–1, υS = 0.157;
and µL = 2, τL = 17 µm–1 and υL = 0.226). In the analysis that follows, the ratio of the
number of small to large particles, nS : nL, is likewise varied, as with the power-law
24
distribution earlier, to examine how changes in the relative concentration of small to

large particles affects the bias between spherical and non-spherical particles. The smaller
the value of nS : nL , the smaller is the relative contribution of small particles to the PSD.
Two types of comparisons between the IOPs of a polydispersion of spheroids are performed
here:
1. A constant aspect ratio is assumed for the whole population and only the slope of the
PSD (ξ or nS : nL) is allowed to vary.
2. The slope of the PSD and the aspect ratio are varied as a function of size following the
observations of Jonasz (1987b) who, utilising scanning electron microscopy, derived the
following shape distribution:
〈G 〉
= 1.28 D 0.22 . (29)
G
The implication of Equation 29 is that the smaller the particles are the more sphere-like
they become. Jonasz (1987b) also found that the larger particles resembled elongated
cylinders with aspect ratios >1. The geometric cross section of an elongated cylinder,
however, is very similar to that of a prolate spheroid and so prolate spheroids are used
here to model larger particles. Thus, the deviation from sphericity of a particle can be
expressed in terms of its aspect ratio, s/t, and diameter of its equal-volume sphere, D,
and Equation 29 becomes:
 
() ()
1 −2
1  s  3
−2 s 3
t
sin −1 1 − s
t  0.22
+  = 1.28 D . (30)
2  t 
()
−2
 1− s 
 t 
Given a size D, this equation is solved to obtain s/t, which is used in the population
model with aspect ratios varying as a function of size (see also Figures 2 and 3 in Jonasz
1987b).
Results for polydispersions

In the following section, the modelled IOPs (c, a and b) of polydispersions of spheroids are
presented. Due to the inability to obtain the VSF of spheroids throughout the size range of interest,
results regarding either the VSF or the backscattering coefficient, bb, are not presented here.
For polydispersions of spheroids, shape effects depend on the relative contributions of small
and large particles to the population and the degree to which particles deviate from a spherical
shape (as indicated by the aspect ratio). In both the power-law and Risoviç (1993) PSD simulations,
with constant and varying aspect ratios, the biases of all the IOPs increase with increasing proportion
of large particles in the population (i.e., as ξ → 3 or as nS : nL → 1012, Figures 13 and 14). This is
a direct consequence of the nearly monotonic change in the bias as a function of size for a
monodispersion (Figure 8). As expected from the results for monodispersions of spheroids, the
biases in attenuation and scattering increase as the aspect ratio departs from one, the absorption
bias also increases with departure from sphericity and with increasing absorption index. In most
cases the biases are >1 (i.e., a spherical model will underestimate a population of spheroids), being
25
2 2
m = 1.05 + i0.01 A m = 1.17 + i0.0001 B
1.8 1.8
1.6 1.6
1.4 1.4
γc
γc
1.2
1.2
1
1
0.8
0.8
1.8
m = 1.05 + i0.01 C 1.4 m = 1.17 + i0.0001 D
1.6
1.3
γa
γa
1.4 1.2
1.1
1.2
1
1
2 2
m = 1.05 + i0.01 E m = 1.17 + i0.0001 F
1.8 1.8
1.6 1.6
1.4 1.4
γb
γb
1.2 1.2
1
1
0.8
0.8
0.6
3.5 3.75 4 4.25 4.5 3.5 3.75 4 4.25 4.5
ξ ξ
Aspect ratio st
x 10 2 1 0.5 0.1
Figure 13 The bias in attenuation, γc (A, B), absorption, γa (C, D), scattering, γb (E, F), and backscattering,
γbb (G, H), for a power-law polydispersion of spheroids relative to a power-law polydispersion of spheres with
the same volume as a function of the power-law exponent, ξ. Each line represents a different aspect ratio, s/t
(legend below the plot). The grey line with dots (legend: ‘x’) denotes the polydispersions of spheroids where
the shape co-varies with size following Jonasz (1983; see text). The dotted vertical lines are used to compare
equivalent size distributions in Figure 14.
<1 only for attenuation and scattering by populations dominated by small particles (i.e., as ξ → 4.5
or as nS : nL → 1017, Figures 13 and 14) and for absorption when small cells dominate and the
aspect ratio is significantly different from one (Figure 13D). Biases of the Risoviç (1993) PSD are
similar to those of the power-law PSD (Figures 13 and 14). Changes in the bias as a function of
the PSD parameter (Figures 13 and 14) are smooth due to the averaging over particles of many
different sizes, as well as to random orientation. The more realistic case of a population varying
in both size and shape exhibits, in general, a larger bias than populations with a constant shape
(Figures 13 and 14) because for all the particles in this population the bias is one or larger while
for those with constant shape the smallest particles very often have biases smaller than one.
26
2.2 A 2.5 B
2 2.2
1.8 1.9
γc
γc
1.6
1.6
1.4
m = 1.05 + i0.01 1.3 m = 1.17 + i0.0001
1.2
1
1
0.7
1.8 1.25
C D
1.2
1.6
1.15
γa
γa
1.4
1.1
m = 1.05 + i0.01 m = 1.17 + i0.0001
1.2
1.05
1 1
2.5 E 2.5 F
2.2 2.2
1.9 1.9
γb
γb
1.6 1.6
1.3 m = 1.05 + i0.01 1.3 m = 1.17 + i0.0001
1 1
0.7 0.7
1013 1014 1015 1016 1017 1013 1014 1015 1016 1017
nS : nL nS : nL
Aspect ratio st
x 10 2 1 0.5 0.1
Figure 14 The biases in attenuation, γc (A, B), absorption, γa (C, D), and scattering, γb (E, F), for a Risoviç
(1993) polydispersion of spheroids relative to a Risoviç polydispersion of spheres with the same volume as
a function of the relative numbers of small to large particles, nS : nL . Each line represents a different aspect
ratio, s/t (legend below the plot). The grey line with dots (legend: ‘x’) denotes the polydispersions of spheroids
where the shape co-varies with size following Jonasz (1983; see text). The dotted vertical lines indicate the
ratios where the size distributions having approximately a power-law slope of ξ = 3.5 and ξ = 4.0, from left
to right respectively, as shown on Figure 13.
The results here are consistent with those of Herring (2002) who investigated the attenuation
bias of a population of particles with a power-law PSD with varying shape as well as varying
refractive index (using the model of the dependence of the index of refraction on size by Zaneveld
et al. 1974). Variations in the attenuation bias were slightly larger in the study by Herring (2002)
than the results obtained here (Figure 13), varying from 2.2 to 1.1 as the power-law slope varied
between 3 ≤ ξ ≤ 5. Varying the imaginary part of the index of refraction between 0.0015 ≤ k ≤
0.01 results in a difference of up to 10% in the attenuation bias, most pronounced for a steeper
PSD (Herring 2002).
27
In general, for the populations of particles modelled here with particle sizes ranging from D =
0.2 to 200 µm, at least 50% of the contribution to c, a and b comes from particles smaller than
D = 10 µm (Figure 15). However, the strongest contribution, as much as 65% (indicated by the
steepness of the curves in the plots), comes from the contribution made by particles of intermediate
sizes belonging to the VDH region. Because in this region the biases are large (Figure 8), models
based on spherical particles will underestimate the IOPs (lines with dots in Figure 15).

0.2 0.5 1 2 5 10 20 50 0.5 1 2 5 10 20 50 200
1 1
A B
0.8 0.8
ci/ Σ cN
Σi=1 ci/ ΣN cN
N
0.6 0.6
0.4 0.4
Σi=1
n
n
0.2 0.2
m = 1.05 + i0.01 m = 1.17 + i0.0001
0 0
1 1
C D
0.8 0.8
ai/ Σ aN
Σi=1 ai/ ΣN aN
N
0.6 0.6 Power-law

0.4 0.4
Σi=1
Risoviç
n
0.2 0.2
m = 1.05 + i0.01 m = 1.17 + i0.0001
0 0
1 1
E F
0.8 0.8
bi/ Σ bN
Σi=1 bi/ ΣN bN
N
0.6 0.6
0.4 0.4
Σi=1
n
0.2 0.2
m = 1.05 + i0.01 m = 1.17 + i0.0001
0 0
0.2 0.5 1 2 5 10 20 50 200 0.2 0.5 1 2 5 10 20 50 200
Aspect ratio st
10 2 1 0.5 0.1
Figure 15 The normalised cumulative contribution to attenuation (c) (A, B), absorption (a) (C, D), and
scattering (b) (E, F), for both the Risoviç (nS : nL = 1016 ) and power-law (ξ = 4) polydispersions. Each line
represents a different aspect ratio, s/t (legend below the plot). The uppermost curves in each panel represent
populations with a power-law size distribution (compare with Figure 13) and the lowermost curves represent
populations with a Risoviç size and shape distribution (compare with Figure 14). In all cases, the greater the
deviation from sphericity (as indicated by the aspect ratio, s/t) the greater is the contribution by larger particles
to the IOPs of the polydispersion.
28
The reader is cautioned that the shapes chosen here, the specific choices for size distribution
(including the size range from Dmin to Dmax), the index of refraction and the internal structure are
all idealisations and may differ from natural environmental conditions. The general trends in the
results, however, are expected to hold true for the natural environment as has been found in many
past studies where idealised optical theory has been used to interpret and invert in situ observations.
In addition, model results presented here provide an order of magnitude estimate for the likely bias
resulting when spherical particles are used to model natural assemblages of particles. Two conclu-
sions can be drawn from the results here and from Herring (2002): (1) biases in IOPs are expected
to be greatest in populations enriched with large particles similar to those in coastal assemblies of
phytoplankton and those in benthic environments and (2) these biases are likely to be smaller than
a factor of three (the biases are likely to be <30% when a population is enriched with small particles
as in the near-surface layer of an oligotrophic ocean).
Observations on the effects of particle shape on IOPs

There are relatively few direct observations on the effects of shape on the IOPs of aquatic particles
(e.g., Volten et al. 1998). Size determination and hence volume are often ambiguous. For example,
when size fractionation is used (i.e., using filters or sieves of increasingly smaller pore sizes)
information on only two of the three dimensions (possibly the largest) of the particles can be
estimated. Similarly, when microscopy is used in sizing, it is difficult to estimate the third (usually
smaller) dimension. Thus, there are uncertainties in both measurements and modelling of IOPs of
non-spherical particles. Several studies provide evidence that aquatic particles in general, and non-
spherical particles in particular, are not optically equivalent to spheres.
Kadyshevich (1977), Voss & Fry (1984) and Volten et al. (1998) measured the VSF and polarised
scattering characteristics of oceanic samples and phytoplankton cultures, respectively, and found
that the polarisation characteristics of scattering are not consistent with spherical particles. Hod-
kinson (1963), Proctor & Barker (1974) and Proctor & Harris (1974) have found that the attenuation
efficiency factor (Qc ) as a function of size of sorted nonspherical particles did not exhibit the
oscillatory behaviour seen for monodispersed spheres and, similar to spheres, attained an asymptotic
value for large particles. This behaviour is similar to Qc for a polydispersion of spheres. However,
the asymptotic value found in those studies was not always two, probably a consequence of the
variety of means by which the ‘size’ of the particles of interest has been determined in the various
studies (Aas 1984).
An example of recent measurements of the near-forward VSF of natural particles from the
Satluj River in India (Figure 16) exhibits two features that are similar to those suggested by the
theoretical analysis presented in this review (Figure 4) and in previous theoretical papers (e.g.,
Kerker 1969). The resonance pattern in the VSF associated with a monodispersion of spheres
disappears for even tightly sorted natural particles, in a similar manner as a polydispersion of
spheres (Figure 16A,C). Associated with a larger cross-sectional area, non-spherical particles
exhibit stronger (diffraction-dominated) forward-peaked VSFs (Figure 16A,C). These results are
consistent with the theoretical results showing no resonance pattern in the VSF of monodispersed
spheroids as a function of angle (Figure 4) and a scattering bias >1 for large particles compared
with wavelength (Figure 8F) due to a higher VSF in the near-forward direction. Similar results
have been recorded by MacCallum et al. (2004) for phytoplankton cultures where the best agreement
in the near-forward VSF was found for spheres with a volume larger by a factor of 1.5.
When Mie-based size inversions are applied to non-spherical and complex particles such as
phytoplankton, unexpected results are produced. For example, inversion of VSF measurements
conducted on a culture of Ceratium longipes suggests a multi-modal population (Figure 17). The
29
104 104
Glass spheres Glass spheres
Sediment Sediment
103 103
β (θ) (unscaled)
β (θ) (unscaled)
102 102
101 101
75–90 µm 100 µm 25–30 µm
100 100
0 1 2 3 4 5 6 0 1 2 3 4 5 6
Scattering angle θ (deg) Satluj River sediment Scattering angle θ (deg)
A B C
Figure 16 Measurements of near-forward scattering. A is a comparison of the shape of the near-forward

scattering between sorted particles (Satluj River sediment; solid line) and glass spheres (line with dots)
75–90 µm in size. An image of the particles used in the measurement of the sediment in A is presented in B.
C is a comparison between sediment (solid line) and glass spheres (line with dots) 25–30 µm in size. Note
the narrowing of the scattering pattern and absence of a secondary maximum for the non-spherical natural
particles (solid lines) compared to polydispersions of spheres sieved similarly (lines with dots). Scattering
measurements were performed with the LISST-100 instrument (data courtesy of Briggs-Whitmire and Agrawal
at Sequoia Scientific, Inc.).
peaks in the volume size distribution, however, are found to correspond to the outer boundary of
the cell, the core part of the cell and the thickness of the appendages (Figure 17).
Summary and future prospect

Together with size, composition and internal structure (not addressed here, however, the reader is
referred to Kitchen et al. 1982, Quirantes & Bernard 2004, 2006), shape has important effects on
IOPs. For a monodispersion of particles, the biases between spheroids and equal-volume spheres
can be larger than a factor of three, while for more realistic polydispersions biases in attenuation,
absorption and scattering are smaller than a factor of three. The size of the particles having the
maximal IOP per unit volume is larger for spheroids than for spheres for all IOPs; this size increases
with non-sphericity.
Many studies have attempted to solve the problem of optical properties of non-spherical particles
by looking for a spherical equivalent. While for certain optical properties and size ranges a single
sphere can provide an adequate model for a non-spherical particle, it has been found that other
properties cannot be modelled with spheres; of these, the degree of polarisation and the VSF in
the backward direction are inherently different for non-spherical geometries (Bohren & Singham
1991). In this survey, following Paramonov (1994b), a polydispersion of spheres with the same
volume and cross-sectional area as a monodispersion of spheroids is used to model the attenuation,
absorption and scattering for the size range in which no ‘exact’ solutions are available (the T-matrix
approach or the ray tracing method). This approximation works well in that it merges into the
T-matrix and ray tracing solutions with very little difference. This method, however, does not
provide an accurate estimate for the VSF or the backscattering coefficient of non-spherical particles.
For the smallest particles, such as viruses in aquatic environments, shape is not likely to affect
the IOPs and hence these particles can be modelled as spheres. Particles with sizes comparable
30
20 µm
1.2
Volume concentration[ µl ]
l
0.8
0.6
0.4
0.2
0
0 50 100 150 200 250
Mean particle diameter (µm)
Figure 17 Volume concentration (assuming spherical particles) inverted from VSF measurements of the
dinoflagellate Ceratium longipes using the Sequoia Scientific LISST-100. The peaks in the size distribution
correspond to different length scales associated with the individual cell.
with the wavelength (e.g., bacteria) have attenuation and scattering biases that are <1. This bias
becomes greater than one as particle size increases (e.g., for microphytoplankton) until it reaches
an asymptote: the ratio of the average cross-sectional area of the spheroid to that of an equal-
volume sphere (Equation 24), which is always >1. For absorption, the bias generally increases with
size until an asymptotic value is reached, as a randomly oriented spheroidal particle is less ‘pack-
aged’ than a sphere.
Backscattering by non-spherical particles is still largely unexplored due to the lack of compu-
tational methods covering much of the range of interest (Figures 9B,D, 10B,D, and 12). The
backscattering bias is, in general, >1 and can be greater by as much as a factor of seven (95% of
the time in Figure 9B) for specific sizes of phytoplankton-like particles. For particles with a very
large absorption coefficient (unrealistic for marine particles), an asymptotic value similar to the
other IOPs is reached (Herring 2002), suggesting that in general, for particles larger than the
wavelength, the backscattering should be more enhanced compared with that of equal-volume
spheres. Despite the complexity observed, it seems sensible to conclude that the backscattering of
spheroids is likely to be significantly larger than that of equal-volume spheres for the sizes relevant
to phytoplankton (Figure 8G,H). In this respect, shape may be a factor contributing to the inability
to account for the bulk backscattering coefficient in the ocean, when spheres are used as a model
for natural particles (e.g., Stramski et al. 2004). Indeed, Morel et al. (2002) used a mixture of
prolate and oblate spheroidal particles (using the T-matrix method) to generate the phase function
31
of small phytoplankton-like particles that was more realistic in the backward directions compared
with that derived from spheres.
For polydispersions of particles with constant or varying shape as a function of size, the biases
in attenuation, absorption and scattering have been found here to be bounded, reaching high values
(270%) only for extreme shapes and size distribution parameters but generally being within about
50% of that of spheres (Figures 13 and 14). While not as large as for monodispersions, these biases
are significant and most often >1, implying that populations of spherical particles perform poorly
as an average, unbiased model.
Diffraction-based instruments provide an opportunity to measure particle size in situ. Given
that measurements are made for angular scattering and that inversions from optical measurements
to obtain particle size are based on Mie theory, shape may cause significant biases for the sizing
of particles. A population of non-spherical particles will appear, on average, larger (and more
dispersed) than a population of equal-volume spheres (Figure 11). In addition, such an inversion
will ‘create’ populations at the tail ends of the size distribution due to the fact that the non-spherical
particles have no resonance pattern in the near-forward scattering as a function of angle (in contrast
to spheres, see Figures 4, 16 and 17; see also Heffels et al. 1996).
Shape is likely to have some effect on optical inversions that are based on Mie theory. In such
inversions, IOPs are used to predict the physical characteristics of the underlying bulk particulate
population. For example, the imaginary part of the index of refraction of phytoplankton has been
found by inverting absorption data using measured size distributions and Mie theory (Bricaud &
Morel 1986). Based on the results of this paper, the inverted k is likely to be an overestimate, with
the bias increasing with increasing phytoplankton size and departure from sphericity. Similarly, an
inversion of the backscattering ratio was used to obtain the real part of the index of refraction for
populations of particles with a power-law size distribution, assuming spherical particles (Tward-
owski et al. 2001, Boss et al. 2004). Results of this work suggest that a spherical model is likely
to underestimate the index of refraction as deviations from sphericity will enhance the backscat-
tering ratio, thus increasing the bias of the inverted index of refraction. Shape effects, on the other
hand, were not found to significantly change the spectral slope of the beam attenuation (Boss et
al. 2001) and thus are not likely to significantly affect the inversion of this parameter to obtain
information on the particulate size distribution. Given the inherent biases associated with using
spheres as models for natural particles, it is sensible to predict that inversions that include non-
spherical characteristics should provide an improvement compared to those based on Mie theory.
This has been the case in several atmospheric studies (e.g., Dubovik et al. 2002, Zhao et al. 2003,
Kocifaj & Horvath 2005).
Shape has important effects on the polarisation of light scattered by marine particles but is a
topic which is beyond the focus of this review. Nevertheless, it is one of the future frontiers in
ocean optics, as currently there is no in situ commercial instrumentation able to measure polarised
scattering. The aquatic community has largely neglected polarisation when studying particulate
suspensions (with a few exceptions, e.g., Quinby-Hunt et al. 2000 and references therein). Studies
by Geller et al. (1985) and Hoovenier et al. (2003) suggest that there is promise in obtaining
information regarding some aspects of particle shape (e.g., departure from sphericity) by analysing
certain elements of the polarised scattering matrix. For example, theoretical shape indices have
been derived based on both linear (Kokhanovsky & Jones 2002) and circular (Hu et al. 2003)
polarisation measurements. In particular, the latter was found to be less sensitive to multiple
scattering. Both were found to be most sensitive at scattering angles in the backward hemisphere.
Polarimetry shows promise especially for extreme shapes and larger particles (Macke & Mishchenko
1996).
Both organic and inorganic aquatic particles are not randomly distributed among shapes but
rather tend to span a limited and non-uniform range of aspect ratios, with spheres being relatively
32
rare. Given the limited amount of data available regarding shape distributions of natural particles,
more measurements of shape parameters are needed; in particular, these are needed as input to
improve inversion models that currently assume spherical particles. Laboratory experiments
designed to measure the effects of shape on optical properties and their consistency with the
predictions presented here and elsewhere are also required so that a more complete picture of the
effect of shape on IOPs can be established.
Acknowledgements
We are indebted to J.R.V. Zaneveld, G. Dall’Olmo and H. Gordon for helpful discussions and
constructive comments on earlier drafts of this manuscript; D. Risoviç for the delight in sharing
the pragmatism of representing particle size distributions; Y.C. Agrawal and A. Briggs-Whitmire
for the scattering measurements and pictures of river sediment; G.R. Fournier for insight into
analytical solutions to ‘the problem’; J.T.O. Kirk for resurrecting the absorption cross section triple
integral that was done on a hand calculator and M.I. Mishchenko for a lifetime of T-matrix code.
This project is supported by the Ocean Optics and Biology programme of the Office of Naval
Research (Contract No. N00014-04-1-0710) to E. Boss and by NASA’s Ocean Biology and Bio-
geochemistry research programme (Contract No. NAG5-12393) to L. Karp-Boss.
References
Aas, E. 1984. Influence of shape and structure on light scattering by marine particles, Report series 53. Institute
of Geophysics, University of Oslo, Oslo, Norway.
Aas, E. 1996. Refractive index of phytoplankton derived from its metabolite composition. Journal of Plankton
Research 18, 2223–2249.
Asano, S. 1979. Light scattering properties of spheroidal particles. Applied Optics 18, 712–723.
Asano, S. & Yamamoto, G. 1975. Light scattering by a spheroidal particle. Applied Optics 14, 29–49.
Aydin, K. 2000. Centimeter and millimeter wave scattering from nonspherical hydrometeors. In Light Scat-
tering by Nonspherical Particles: Theory, Measurements, and Applications, M.I. Mishchenko et al.
(eds). San Diego, California: Academic Press, 451–479.
Baba, J. & Komar, P.D. 1981. Settling velocities of irregular grains at low Reynolds numbers. Journal of
Sedimentary Petrology 51, 121–128.
Bickmore, B.R., Nagy, K.L., Sandlin, P.E. & Crater, T.S. 2002. Quantifying surface areas of clays by atomic
force microscopy. American Mineralogist 87, 780–783.
Bohren, C.F. & Huffman, D.R. 1983. Absorption and Scattering of Light by Small Particles. New York: Wiley.
Bohren, C.F. & Singham, S.B. 1991. Backscattering by nonspherical particles: a review of methods and
suggested new approaches. Journal of Geophysical Research 96, 5269–5277.
Boss, E., Stramski, D., Bergmann, T., Pegau, W.S. & Lewis, M. 2004. Why should we measure optical
backscattering? Oceanography 17, 44–49.
Boss, E., Twardowski, M.S. & Herring, S. 2001. The shape of the particulate beam attenuation spectrum and
its relation to the size distribution of oceanic particles. Applied Optics 40, 4885–4893.
Bricaud, A. & Morel, A. 1986. Light attenuation and scattering by phytoplankton cells: a theoretical modeling.
Applied Optics 25, 571–580.
Carder, K.L., Beardsley, G.F. & Pak, H. 1971. Particle size distributions in the eastern equatorial Pacific.
Journal of Geophysical Research 76, 5070–5077.
Carder, K.L., Betzer, P.R. & Eggiman, D.W. 1974. Physical, chemical and optical measures of suspended
particle concentrations: Their intercomparison and application to the west African shelf. In Suspended
Solids in Water, R.J. Gibbs (ed.). New York: Plenum, 173–193.
33
Cauchy, A.L. 1832. Mémoire sur la rectification des courbes et la quadrature des surfaces courbes. Paris:
Académie des Sciences.
Chýlek, P., Grams, G.W. & Pinnick, R.G. 1976. Light scattering by irregular randomly oriented particles.
Science 193, 480–482.
Dubovik, O., Holben, B.N., Lapyonok, T., Sinyuk, A., Mishchenko, M.I., Yang, P. & Slutsker, I. 2002. Non-
spherical aerosol retrieval method employing light scattering by spheroids. Geophysical Research
Letters 29, 1415–1418.
Duysens, L.M.N. 1956. The flattening of the absorption spectrum of suspensions, as compared to that of
solutions. Biochimica et Biophysica Acta 19, 1–12.
Fournier, G.R. & Evans, B.T.N. 1991. Approximation to extinction efficiency for randomly oriented spheroids.
Geller, P.E., Tsuei, T.G. & Barber, P.W. 1985. Information content of the scattering matrix for spheroidal
particles. Applied Optics 24, 2391–2396.
Gordon, H.R. 2006. Backscattering of light from disklike particles: is fine-scale structure or gross morphology
more important? Applied Optics 45, 7166–7173.
Gordon, H.R. & Du, T. 2001. Light scattering by nonspherical particles: application to coccoliths detached
from Emiliania huxleyi. Limnology and Oceanography 46, 1438–1454.
Heffels, C.M.G., Verheijen, P.J.T., Heitzmann, D. & Scarlett, B. 1996. Correction of the effect of particle
shape on the size distribution measured with a laser diffraction instrument. Particle and Particle
Systems Characterization 13, 271–279.
Herring, S.G. 2002. A systematic survey of the modeled optical properties of nonspherical marine-like particles.
MS thesis, Oregon State University, Corvallis, Oregon.
Hillebrand, H., Dürselen, C.-D., Kirschtel, D., Pollingher, U. & Zohary, T. 1999. Biovolume calculation for
pelagic and benthic microalgae. Journal of Phycology 35, 403–424.
Hodkinson, J.R. 1963. Light scattering and extinction by irregular particles larger than the wavelength. In
Electromagnetic Scattering, M. Kerker (ed.). New York: Macmillan, 87–100.
Hoovenier, J.W., Volten, H., Muñoz, O., van der Zande, W.J. & Waters, L.B.F.M. 2003. Laboratory studies of
scattering matrices for randomly oriented particles: potentials, problems, and perspectives. Journal
of Quantitative Spectroscopy and Radiative Transfer 79–80, 741–755.
Hu, Y.-X., Yang, P., Lin, B., Gibson, G. & Hostetler, C. 2003. Discriminating between spherical and non-
spherical scatterers with lidar using circular polarization: a theoretical study. Journal of Quantitative
Spectroscopy and Radiative Transfer 79–80, 757–764.
Jerlov, N.G. 1968. Optical Oceanography. Amsterdam: Elsevier.
Jerlov, N.G. 1976. Marine Optics. Amsterdam: Elsevier Scientific Publishing Company, Elsevier Oceanogra-
phy Series 14. 2nd edition.
Jonasz, M. 1983. Particle-size distributions in the Baltic. Tellus 35B, 346–358.
Jonasz, M. 1987a. Nonspherical sediment particles: comparison of size and volume distributions obtained
with an optical and resistive particle counter. Marine Geology 78, 137–142.
Jonasz, M. 1987b. Nonsphericity of suspended marine particles and its influence on light scattering. Limnology
and Oceanography 32, 1059–1065.
Jonasz, M. 1991. Size, shape, composition, and structure of microparticles from light scattering. In Principles,
Methods, and Application of Particle Size Analysis, J.P.M. Syvitski (ed.). Cambridge: Cambridge
University Press, 143–162.
Jonasz, M. & Fournier, G.F. 1996. Approximation of the size distribution of marine particles by a sum of log-
normal functions. Limnology and Oceanography 41, 744–754. Errata published 1999, Limnology and
Oceanography 44, 1358.
Jonasz, M. & Fournier, G. 2007. Light Scattering by Particles in Water: Theoretical and Experimental
Foundations. Amsterdam: Elsevier Science.
Junge, C.E. 1963. Air Chemistry and Radioactivity. New York: Academic Press.
Kadyshevich, Ye. A. 1977. Light-scattering matrices of inshore waters of the Baltic Sea. Izvestiya, Atmospheric
and Oceanic Physics 13, 77–78. Translated by the American Geophysical Union from Izvestiia
Akademii nauk SSSR. Fizika atmosfery i okeana.
34
Karp-Boss, L. & Jumars, P.A. 1998. Motion of diatom chains in steady shear flow. Limnology and Oceanog-
raphy 43, 1767–1773.
Kerker, M. 1969. The Scattering of Light and Other Electromagnetic Radiation. San Diego, California:
Academic Press.
Kirk, J.T.O. 1976. A theoretical analysis of the contribution of algal cells to the attenuation of light within
natural waters. III. Cylindrical and spheroidal cells. New Phytologist 77, 341–358.
Kirk, J.T.O. 1994. Light and Photosynthesis in Aquatic Environments. Cambridge: Cambridge University
Press. 2nd edition.
Kitchen, J.C., Zaneveld, J.R.V. & Pak, H. 1982. Effect of particle size distribution and chlorophyll content
on bean attenuation spectra. Applied Optics 21, 3913–3918.
Kocifaj, M. & Horvath, H. 2005. Retrieval of size distribution for urban aerosols using multispectral optical
data. Journal of Physics: Conference Series 6, 97–102.
Kokhanovsky, A.A. 2003. Optical properties of irregularly shaped particles. Journal of Physics D: Applied
Physics 36, 915–923.
Kokhanovsky, A.A. & Jones, A.R. 2002. The cross-polarization of light by large non-spherical particles.
Journal of Physics D: Applied Physics 35, 1903–1906.
Kokhanovsky, A.A. & Zege, E.P. 1997. Optical properties of aerosol particles: a review of approximate
analytical solutions. Journal of Aerosol Science 28, 1–21.
Komar, P.D. & Reimers, C.E. 1978. Grain shape effects on settling rates. Journal of Geology 86, 193–209.
Latimer, P., Brunsting, A., Pyle, B.E. & Moore, C. 1978. Effects of asphericity on single particle scattering.
MacCallum, I., Cunningham, A. & McKee, D. 2004. The measurement and modelling of light scattering by
phytoplankton cells at narrow forward angles. Journal of Optics A: Pure and Applied Optics 6, 698–702.
Macke, A. & Mishchenko, M.I. 1996. Applicability of regular particle shapes in light scattering calculations
for atmospheric ice crystals. Applied Optics 35, 4291–4296.
Macke, A., Mishchenko, M.I., Muinonen, K. & Carlson, B.E. 1995. Scattering of light by large nonspherical
particles: ray-tracing approximation versus T-matrix method. Optics Letters 20, 1934–1936.
Mie, G. 1908. Beiträge zur Optik trüber Medien, speziell kolloidaler Metallösungen. Annalen der Physik 25,
377–445.
Mishchenko, M.I., Hoovenier, J.W. & Travis, L.D. (eds). 2000. Light Scattering by Nonspherical Particles:
Theory, Measurements, and Applications. San Diego, California: Academic Press.
Mishchenko, M.I., Travis, L.D. & Lacis, A.A. 2002. Scattering, Absorption and Emission of Light by Small
Particles. New York: Cambridge University Press.
Mobley, C.D. 1994. Light and Water: Radiative Transfer in Natural Waters. San Diego, California: Academic Press.
Morel, A. 1973. Diffusion de la lumière par les eaux de mer: résultats expérimentaux et approche theorique.
AGARD Lecture Series 61, 3.1.1–3.1.76.
Morel, A., Antoine, D. & Gentili, B. 2002. Bidirectional reflectance of oceanic waters: accounting for Raman
emission and varying particle scattering phase function. Applied Optics 41, 6289–6306.
Morel, A. & Bricaud, A. 1981. Theoretical results concerning light absorption in a discrete medium, and
application to specific absorption of phytoplankton. Deep-Sea Research 28A, 1375–1393.
Paramonov, L.E. 1994a. A simple formula for estimation of the absorption cross section of biological suspen-
sions. Optics and Spectroscopy 77, 506–512. Translated from Optika I Spektroskopiya.
Paramonov, L.E. 1994b. On optical equivalence of randomly oriented ellipsoidal and polydisperse spherical
particles. The extinction, scattering and absorption cross sections. Optics and Spectroscopy 77, 589–592.
Translated from Optika i Spektroskopiya.
Pozdnyakov, D. & Grassl, H. 2003. Colour of Inland and Coastal Waters: A Methodology for Its Interpretation.
Chichester, U.K.: Praxis.
Proctor, T.D. & Barker, D. 1974. The turbidity of suspensions of irregularly shaped diamond particles. Journal
of Aerosol Science 5, 91–99.
Proctor, T.D. & Harris, G.W. 1974. The turbidity of suspensions of irregular quartz particles. Journal of Aerosol
Science 5, 81–90.
35
Quinby-Hunt, M.S., Hull, P.G. & Hunt, A.J. 2000. Polarized light scattering in the marine environment. In
Light Scattering by Nonspherical Particles: Theory, Measurements, and Applications, M.I. Mish-
chenko et al. (eds). San Diego, California: Academic Press, 525–554.
Quirantes, A. & Bernard, S. 2004. Light scattering by marine algae: two-layer spherical and nonspherical
models. Journal of Quantitative Spectroscopy and Radiative Transfer 89, 311–321.
Quirantes, A. & Bernard, S. 2006. Light-scattering methods for modelling algal particles as a collection of
coated and/or nonspherical particles. Journal of Quantitative Spectroscopy and Radiative Transfer
100, 315–324.
Risoviç, D. 1993. Two-component model of sea particle size distribution. Deep-Sea Research I 40, 1459–1473.
Roesler, C.S. & Boss, E. 2007. In situ measurement of the inherent optical properties (IOPs) and potential
for harmful algal bloom detection and coastal ecosystem observations. In Real-Time Coastal Observing
Systems for Ecosystem Dynamics and Harmful Algal Blooms, M. Babin et al. (eds). Paris: UNESCO
Publishing, in press.
Sheldon, R.W., Prakash, A. & Sutcliffe, W.H. Jr. 1972. The size distribution of particles in the ocean. Limnology
Shepelevich, N.V., Prostakova, I.V. & Lopatin, V.N. 2001. Light-scattering by optically soft randomly oriented
spheroids. Journal of Quantitative Spectroscopy and Radiative Transfer 70, 375–381.
Shifrin, K.S. 1988. Physical Optics of Ocean Water. New York: American Institute of Physics, AIP translation
series. Originally published as Vvedenie v optiku morya 1983.
Siegel, D.A. 1998. Resource competition in a discrete environment: Why are plankton distributions paradox-
ical? Limnology and Oceanography 43, 1133–1146.
Stramski, D., Boss, E., Bogucki, D. & Voss, K.J. 2004. The role of seawater constituents in light backscattering
in the ocean. Progress in Oceanography 61, 27–56.
Stramski, D., Bricaud, A. & Morel, A. 2001. Modeling the inherent optical properties of the ocean based on
the detailed comparison of the planktonic community. Applied Optics 40, 2929–2945.
Stramski, D. & Kiefer, D.A. 1991. Light scattering by microorganisms in the open ocean. Progress in
Oceanography 28, 343–383.
Syvitski, J.P.M., Asprey, K.W. & Leblanc, K.W.G. 1995. In-situ characteristics of particles settling within a
deep-water estuary. Deep-Sea Research Part II: Topical Studies in Oceanography 42, 223–256.
Twardowski, M.S., Boss, E., MacDonald, J.B., Pegau, W.S., Barnard, A.H. & Zaneveld, J.R. 2001. A model
for estimating bulk refractive index from the optical backscattering ratio and the implications for
understanding particle composition in Case I and Case II waters. Journal of Geophysical Research
106, 14129–14142.
Twardowski, M.S., Lewis, M.R., Barnard, A.H. & Zaneveld, J.R. 2005. In-water instrumentation and platforms
for ocean color remote sensing applications. In Remote Sensing of Coastal Aquatic Environments:
Technologies, Techniques and Applications, R.L. Miller et al. (eds). Dordrecht: Springer, 69–100.
van de Hulst, H.C. 1981. Light Scattering by Small Particles. New York: Dover.
Volten, H., de Haan, J.F., Hoovenier, J.W., Schreurs, R. & Vassen, W. 1998. Laboratory measurements of
angular distributions of light scattered by phytoplankton and silt. Limnology and Oceanography 43,
1180–1197.
Voss, K.J. & Fry, E.S. 1984. Measurement of the Mueller matrix for ocean water. Applied Optics 23,
4427–4439.
Waterman, P.C. 1971. Symmetry, unitarity, and geometry in electromagnetic scattering. Physical Review D 3,
825–839.
Zaneveld, J.R.V., Roach, D.M. & Pak, H. 1974. The determination of the index of refraction distribution of
oceanic particles. Journal of Geophysical Research 79, 4091–4095.
Zhao, T.X.-P., Laszlo, I., Dubovik, O., Holben, B.N., Sapper, J., Tanré, D. & Pietras, C. 2003. A study of the
effect of non-spherical dust particles on the AVHRR aerosol optical thickness retrievals. Geophysical
Research Letters 30, 1317–1320.
36
APPENDIX: NOTATION
Notation used following Mobley (1994) closely. The actual units used are given in the text, however,
only the dimensions are provided in this table for mass, M, length, L, and time, T, or angular
measure as indicated.
Symbol Definition Dimension

a Absorption coefficient L–1
b Scattering coefficient L–1
b̃ Backscattering ratio dimensionless
bb Backscattering coefficient L–1
Ca Absorption cross section of a particle L2
Cb Scattering cross section of a particle L2
C bb Backscattering cross section of a particle L2
Cc Attenuation cross section of a particle L2
c Attenuation coefficient L–1
D Particle size represented by diameter of an equal-volume sphere L
D0 Reference diameter of size range L
E(0) Irradiance at the light source M L–1 T –3
E(R) Irradiance at distance R from the light source M L–1 T –3
f (D) Particulate size distribution # L–4
G Geometrical cross-sectional area of a sphere L2
〈G〉 Average geometrical cross-sectional area of a non-sphere L2
I Radiant intensity M L–1 T –3 sr –1
k Imaginary part of the relative index of refraction dimensionless
k Wave number of the incident light L–1
m Complex relative index of refraction dimensionless
N Number of particles per unit volume # L–3
n Real part of the relative index of refraction dimensionless
n0 Number concentration of particles at the reference diameter D0 # L–4
nS Number concentration of small particles # L–4
nL Number concentration of large particles # L–4
Qa Absorption efficiency factor of a particle dimensionless
Qb Scattering efficiency factor of a particle dimensionless
Qbb Backscattering efficiency factor of a particle dimensionless
Qc Attenuation efficiency factor of a particle dimensionless
R Arbitrary path length of light L
s Rotational axis of a spheroid L
s/t Aspect ratio of a spheroid dimensionless
t Equatorial axis of a spheroid L
V Particle volume L3
x Size parameter dimensionless
α* Specific absorption coefficient of a particle L–1
αa Volume-normalised absorption cross section L–1
αb Volume-normalised scattering cross section L–1
αbb Volume-normalised backscattering cross section L–1
αc Volume-normalised attenuation cross section L–1
β(θ) Volume scattering function (VSF) L–1sr –1
β(θ)
˜ Volume scattering phase function sr –1
γa Absorption bias dimensionless
γb Scattering bias dimensionless
37
Symbol Definition Dimension

γbb Backscattering bias dimensionless
γc Attenuation bias dimensionless
θ Scattering angle radians (rad)
λ Wavelength of the incident light L
µS Small-particle generalised gamma distribution parameter dimensionless
µL Large-particle generalised gamma distribution parameter dimensionless
ξ Slope of the power-law size distribution dimensionless
ρ Phase shift parameter dimensionless
τS Small-particle generalised gamma distribution parameter L–1
τL Large-particle generalised gamma distribution parameter L–1
υS Small-particle generalised gamma distribution parameter dimensionless
υL Large-particle generalised gamma distribution parameter dimensionless
ϕ Azimuth angle radians (rad)
Ψ Angular direction into which light is scattered radians (rad)
Ω Solid angle into which light is scattered steradians (sr)
38
50931_book.fm Page 39 Tuesday, May 1, 2007 4:43 PM
Oceanography and Marine Biology: An Annual Review, 2007, 45, 39-88

© R. N. Gibson, R. J. A. Atkinson, and J. D. M. Gordon, Editors
Taylor & Francis
GLOBAL ECOLOGY OF THE GIANT KELP MACROCYSTIS:

FROM ECOTYPES TO ECOSYSTEMS
MICHAEL H. GRAHAM1,2, JULIO A. VÁSQUEZ2,3 & ALEJANDRO H. BUSCHMANN4
1Moss Landing Marine Laboratories, 8272 Moss Landing Road,
Moss Landing, California 95039, U.S.

E-mail: mgraham@mlml.calstate.edu
2Centro de Estudios Avanzados de Zonas Aridas (CEAZA - www.ceaza.cl)
3Departamento Biología Marina, Facultad de Ciencias del Mar,
Universidad Católica del Norte, Coquimbo, Chile

4Centro de Investigación y Desarrollo en Ambientes y Recursos Costeros (i~mar),
Universidad de Los Lagos, Casilla 557, Puerto Montt, Chile
Abstract The giant kelp Macrocystis is the world’s largest benthic organism and most widely
distributed kelp taxon, serving as the foundation for diverse and energy-rich habitats that are of
great ecological and economical importance. Although the basic and applied literature on Macro-
cystis is extensive and multinational, studies of large Macrocystis forests in the northeastern Pacific
have received the greatest attention. This review synthesises the existing Macrocystis literature into
a more global perspective. During the last 20 yr, the primary literature has shifted from descriptive
and experimental studies of local Macrocystis distribution, abundance and population and commu-
nity structure (e.g., competition and herbivory) to comprehensive investigations of Macrocystis life
history, dispersal, recruitment, physiology and broad-scale variability in population and community
processes. Ample evidence now suggests that the genus is monospecific. Due to its highly variable
physiology and life history, Macrocystis occupies a wide variety of environments (intertidal to
60+ m, boreal to warm temperate) and sporophytes take on a variety of morphological forms.
Macrocystis sporophytes are highly responsive to environmental variability, resulting in differential
population dynamics and effects of Macrocystis on its local environment. Within the large subtidal
giant kelp forests of southern California, Macrocystis sporophytes live long, form extensive surface
canopies that shade the substratum and dampen currents, and produce and retain copious amounts
of reproductive propagules. The majority of subtidal Macrocystis populations worldwide, however,
are small, narrow, fringing forests that are productive and modify environmental resources (e.g.,
light), yet are more dynamic than their large southern California counterparts with local recruitment
probably resulting from remote propagule production. When intertidal, Macrocystis populations
exhibit vegetative propagation. Growth of high-latitude Macrocystis sporophytes is seasonal, coin-
cident with temporal variability in insolation, whereas growth at low latitudes tracks more episodic
variability in nutrient delivery. Although Macrocystis habitat and energy provision varies with
such ecotypic variability in morphology and productivity, the few available studies indicate that
Macrocystis-associated communities are universally diverse and productive. Furthermore, temporal
and spatial variability in the structure and dynamics of these systems appears to be driven by
processes that regulate Macrocystis distribution, abundance and productivity, rather than the con-
sumptive processes that make some other kelp systems vulnerable to overexploitation. This global
synthesis suggests that the great plasticity in Macrocystis form and function is a key determinant
of the great global ecological success of Macrocystis.
39
MICHAEL H. GRAHAM, JULIO A. VÁSQUEZ & ALEJANDRO H. BUSCHMANN
Introduction
Kelp beds and forests represent some of the most conspicuous and well-studied marine habitats.
As might be expected, these diverse and productive systems derive most of their habitat structure
and available energy (fixed carbon) from the kelps, a relatively diverse order of large brown algae
(Laminariales, Phaeophyceae; ~100 species). Kelps and their associated communities are conspic-
uous features of temperate coasts worldwide (Lüning 1990), including all of the continents except
Antarctica (Moe & Silva 1977), and the proximity of such species-rich marine systems to large
coastal human populations has subsequently resulted in substantial extractive and non-extractive
industries (e.g., Leet et al. 2001). It is therefore not surprising that the basic and applied scientific
literature on kelps is extensive.
Our present understanding of the ecology of kelp taxa is not uniform, as the giant kelp Macro-
cystis has received the greatest attention. Macrocystis is the most widely distributed kelp genus in
the world, forming dense forests in both the Northern and Southern hemispheres (Figure 1). The
floating canopies of Macrocystis adult sporophytes also have great structural complexity and high
rates of primary productivity (Mann 1973, Towle & Pearse 1973, Jackson 1977, North 1994).
Furthermore, although Macrocystis primary production can fuel secondary productivity through
direct grazing, most fixed carbon probably enters the food web through detrital pathways or is
exported from the system (e.g., Gerard 1976, Pearse & Hines 1976, Castilla & Moreno 1982,
Castilla 1985, Inglis 1989, Harrold et al. 1998, Graham 2004). In some regions, such habitat and
energy provision can support from 40 to over 275 common species (Beckley & Branch 1992,
Vásquez et al. 2001, Graham 2004).
Venerated by Darwin (1839), the ecological importance of Macrocystis has long been recogn-
ised. The genus, however, did not receive thorough ecological attention until the 1960s when various
Macrocystis research programmes began in California, and later in British Columbia, Chile, México,
and elsewhere. Since that time, several books and reviews and hundreds of research papers have
appeared in both the primary and secondary literature, primarily emphasising the physical and biotic
factors that regulate Macrocystis distribution and abundance, recruitment, reproductive strategies
and the structure and organisation of Macrocystis communities (see reviews by North & Hubbs
1968, North 1971, 1994, Dayton 1985a, Foster & Schiel 1985, North et al. 1986, Vásquez &
Buschmann 1997).
This review synthesises this rich literature into a global perspective of Macrocystis ecology
and such a review is timely for three reasons. First, the last review of Macrocystis ecology was
done by North (1994) and thoroughly covered the literature until 1990, yet there has been significant
progress on many aspects of Macrocystis ecology since that time. Second, during the last 15–20 yr
the general focus of Macrocystis research (and that of kelps in general) has shifted from descriptive
and experimental studies of local Macrocystis distribution, abundance and population and commu-
nity structure (e.g., competition and herbivory) to comprehensive investigations of Macrocystis life
history, dispersal, recruitment, physiology and broad-scale variability in population and community
processes. Finally, previous reviews of Macrocystis ecology have been from an inherently regional
perspective (e.g., California or Chile) and there is currently no truly global synthesis. This last
aspect is of great concern because it effectively partitions kelp forest researchers into provincial
programmes and limits cross-fertilisation of ideas. Such a limitation is compounded by the great
worldwide scientific and economic importance of this genus, the acclimatisation of Macrocystis to
regional environments, and the recent finding that gene flow occurs among the most geographically
distant regions over ecological timescales (Coyer et al. 2001). Therefore, the goal here is not to review
the existing Macrocystis literature in its entirety, but rather to (1) focus on progress made during
the last 15 yr, (2) discuss the achievements of Macrocystis research programmes worldwide and
(3) identify deficiencies in the understanding of Macrocystis ecology that warrant future investigation.
40
Alaska
British
Columbia Washington
Oregon
California
Baja California
Mexico
41
Peru
South Africa
Chile Tristan de Cunha Is. South Australia New Zealand

Chatham Is.
Argentina Gough Is. Crozet Is. Amsterdam/St.Paul Is.
Tasmania Bounty Is.
Prince Edward Is. Kerguelen Is.
Falkland Is. Auckland Is.
Heard Is. Antipodes Is.
South Georgia Is.
Campbell Is.
Figure 1 Global distribution of the giant kelp Macrocystis. Locations are given for distinct Macrocystis mainland and island populations determined directly from
citations herein.
GLOBAL ECOLOGY OF THE GIANT KELP MACROCYSTIS: FROM ECOTYPES TO ECOSYSTEMS
In particular, it is now recognised that great variability exists in Macrocystis morphology, physi-
ology, population dynamics and community interactions at the global scale and it is considered
that such ecotypic variability is key to understanding the role of Macrocystis in kelp systems
worldwide.
Organismal biology of Macrocystis

Most of the biological processes that ultimately prove to be important in regulating the dynamics
and structure of Macrocystis populations and communities (e.g., morphological complexity, photo-
synthesis, growth, reproductive output, gene flow) operate primarily at the scale of individual
organisms. The standard means of studying Macrocystis organismal biology continues to be through
laboratory studies. Clearly, laboratory studies allow researchers to address various processes under
controlled environmental conditions, but in many cases the reliance on laboratory studies has been
due to technical limitations in collecting organismal data in situ. Various technological advances
since the 1960s (most occurring in the last two decades), however, have resulted in a surge of
studies of Macrocystis evolutionary history, distribution, life history, growth, productivity and
reproduction.
Evolutionary history
The order Laminariales has traditionally included five families (Chordaceae, Pseudochordaceae,
Alariaceae, Laminariaceae, Lessoniaceae) but various ultrastructural and molecular data suggest
that subordinal classification (i.e., families, genera, and species) is in need of significant revision
(Druehl et al. 1997, Yoon et al. 2001, Lane et al. 2006). For example, the Chordaceae and
Pseudochordaceae should not be included in the Laminariales (Saunders & Druehl 1992, 1993,
Druehl et al. 1997) and a new family has been proposed (Costariaceae; Lane et al. 2006). The order
is presumed to have originated in the northeast Pacific (Estes & Steinberg 1988, Lüning 1990) and
molecular studies have estimated the date of origin to be between 15 and 35 million yr ago
(Saunders & Druehl 1992). Within the order, the genus Macrocystis was formerly assigned to the
family Lessoniaceae (including Lessonia, Lessoniopsis, Dictyoneurum, Dictyoneuropsis, Nereo-
cystis, Postelsia and Pelagophycus; Setchell & Gardner 1925), which was considered paraphyletic
to the Laminariaceae (Druehl et al. 1997, Yoon et al. 2001). Recent molecular studies, however,
have found that Lessonia, Lessoniopsis, Dictyoneurum and Dictyoneuropsis are actually in phylo-
genetic clades that do not include Macrocystis, and that Macrocystis, Nereocystis, Postelsia and
Pelagophycus group together in a derived clade that is nested well within the Laminariaceae (Lane
et al. 2006), with Pelagophycus porra being the most closely related taxon to Macrocystis.
Species classification within the genus Macrocystis was originally based on blade morphology
yielding over 17 species (see review by North (1971)). Blade morphology was then considered a
plastic trait strongly affected by environmental conditions and subsequently all 17 Macrocystis
species were synonymised with Macrocystis pyrifera (Hooker 1847). Macrocystis species were
later described based on holdfast morphology ultimately leading to the current recognition of three
species: M. pyrifera (conical holdfast; Figure 2A), M. integrifolia (rhizomatous holdfast; Figure 2B),
and M. angustifolia (mounding rhizomatous holdfast) (Howe 1914, Setchell 1932, Womersley
1954, Neushul 1971). The fourth currently recognised species, M. laevis, was described by Hay
(1986), again based on blade morphology (M. laevis has smooth fleshy blades and a M. pyrifera-
type conical holdfast). Four lines of evidence, however, suggest that this current classification of
Macrocystis is also in need of revision: (1) M. pyrifera, M. integrifolia and M. angustifolia are
interfertile (Lewis et al. 1986, Lewis & Neushul 1994; interfertility with M. laevis has not been
42
A B
C D
Figure 2 Macrocystis holdfast morphologies and sporophyte spacing. (A) Holdfast of pyrifera-form sporo-
phyte from La Jolla, southern California. (Published with permission of Scott Rumsey.) (B) Holdfast of
integrifolia-form sporophyte from Huasco, northern Chile. (Photograph by Michael Graham.) (C) Vertical
structure of pyrifera-form population from San Clemente Island (15 m depth), southern California; note average
sporophyte spacing is 3–7 m. (Published with permission of Enric Sala.) (D) Vertical structure of angustifolia-
form population from Soberanes Point (3 m depth), central California; note average sporophyte spacing is
10–50 cm. (Published with permission of Aurora Alifano.)
tested); (2) intermediate morphologies have been observed in the field (Setchell 1932, Neushul
1959, Womersley 1987, Brostoff 1988); (3) in addition to blade morphology (Hurd et al. 1997),
holdfast morphology is phenotypically plastic (Setchell 1932, M.H. Graham, unpublished data);
and most importantly, (4) patterns of genetic relatedness among all four species are not in concor-
dance with current morphological classification (Coyer et al. 2001). This evidence strongly supports
the recognition of the genus Macrocystis as a single morphologically plastic species, with global
populations linked by non-trivial gene flow. For the purpose of this review, therefore, the four
currently recognised species are referred to simply as giant kelp, Macrocystis.
Biogeographic studies of extant kelp in the north Pacific suggest that the bi-hemispheric
(antitropical) global distribution of Macrocystis developed as the genus arose in the Northern
Hemisphere and subsequently colonised the Southern Hemisphere (North 1971, Nicholson 1978,
Estes & Steinberg 1988, Lüning 1990, Lindberg 1991). Alternatively, North (1971) and Chin et al.
(1991) proposed a Southern Hemisphere origin of the genus, the latter via vicariant processes that
have been questioned (Lindberg 1991). Recently, Coyer et al. (2001) studied the global phylogeog-
raphy of Macrocystis using recombinant DNA internal transcribed spacer (ITS1 and ITS2) regions.
In addition to suggesting that the morphological species description of M. pyrifera, M. integrifolia,
M. angustifolia and M. laevis has no systematic support, Coyer et al. (2001) described a well-
resolved phylogeographic pattern in which Southern Hemisphere Macrocystis populations nested
within Northern Hemisphere populations, linked by Macrocystis populations on the Baja California
Peninsula, Mexico. This pattern, and the greater genetic diversity among Macrocystis populations
in the Northern Hemisphere (within-region sequence divergences 1.7% and 1.2% for ITS1 and
ITS2, respectively) relative to their Southern Hemisphere counterparts (within-region sequence
43
divergences 0.8% and 0.6% for ITS1 and ITS2, respectively), supports a northern origin of the
genus with subsequent range expansion to include the Southern Hemisphere (Coyer et al. 2001);
Coyer et al. (2001) suggested that gene flow across the equator may have occurred as recently as
10,000 yr ago.
Despite such progress, however, many questions remain regarding the evolutionary history of
Macrocystis. Most importantly, how can this single, globally distributed species maintain gene flow
throughout its range, yet at a regional scale exhibit relatively high geographic uniformity in such
seemingly important characters as blade and holdfast morphology (i.e., ecotypes or forms)? The
data of Coyer et al. (2001) suggest that simple founder effects may have resulted in the unique
morphologies of the laevis form at the Prince Edward Islands (including Marion Island) and
angustifolia form in Australia. The smooth-bladed laevis form has been found occasionally at the
Falkland Islands (van Tüssenbroek 1989a) and a recent description from Chiloé Island, Chile
(Aguilar-Rosas et al. 2003), is probably a misidentification of sporophylls as vegetative blades
(Gutierrez et al. 2006). Still, despite the apparently high gene flow and morphological plasticity,
the distinct forms with distinct ecologies can dominate different habitats often adjacent to each
other (e.g., integrifolia form in shallow water vs. pyrifera form in deep water). The identification
of which Macrocystis form is present within a region will aid in the understanding of the region’s
ecology (see ‘Population’ section, p. 54). In this context, it is hypothesised that the great plasticity
in Macrocystis form and function may, in fact, be an adaptive trait resulting in its great global
ecological success. Studies testing this hypothesis will require a better understanding of the nature
of Macrocystis morphological plasticity, including biomechanics, structural biochemistry and quan-
titative genetics studies of genes regulating Macrocystis form.
Distribution
Macrocystis distributional patterns have been well described (especially in the Northern Hemisphere)
due primarily to the large stature of Macrocystis sporophytes and ability to sense their surface canopies
remotely from aircraft or satellites (Jensen et al. 1980, Hernández-Carmona et al. 1989a,b, 1991,
Augenstein et al. 1991, Belsher and Mouchot 1992, Deysher 1993, North et al. 1993, Donnellan
2004). Macrocystis typically grows on rocky substrata between the low intertidal and ~25 m depth
(Figure 3; Rigg 1913, Crandall 1915, Baardseth 1941, Papenfuss 1942, Scagel 1947, Guiler 1952,
1960, Cribb 1954, Chamberlain 1965, Neushul 1971, Foster & Schiel 1985, Westermeier & Möller
1990, van Tüssenbroek 1993, Schiel et al. 1995, Graham 1997, Spalding et al. 2003, Vega et al.
2005) and is distributed in the northeast Pacific from Alaska to México, along the west and southeast
coasts of South America from Perú to Argentina, in isolated regions of South Africa, Australia and
New Zealand and around most of the sub-Antarctic islands to 60°S (Figure 1; Crandall 1915,
Baardseth 1941, Cribb 1954, Papenfuss 1964, Chamberlain 1965, Neushul 1971, Hay 1986, Ste-
genga et al. 1997). In unique circumstances, sexually reproducing populations can exist in deep
water (50–60 m; Neushul 1971 (Argentina), Perissinotto & McQuaid 1992 (Prince Edward Islands)),
in sandy habitats (Neushul 1971) and unattached populations that reproduce vegetatively can exist
in the water column (North 1971) or shallow basins (Moore 1943, Gerard & Kirkmann 1984, van
Tüssenbroek 1989b). High latitudinal limits appear to be set by increased wave action (Foster &
Schiel 1985, Graham 1997) and decreased insolation (Arnold & Manley 1985, Jackson 1987),
whereas low latitudinal limits appear to be set by low nutrients associated with warmer (non-upwelling)
waters (Ladah et al. 1999, Hernández-Carmona et al. 2000, 2001, Edwards 2004) or competition with
warm-tolerant species (e.g., Eisenia arborea on the Baja California Peninsula, Mexico; Edwards
& Hernández-Carmona 2005). The upper shallow limits of Macrocystis populations are ultimately
regulated by the increased desiccation and high ultraviolet and/or photosynthetically active radiation
(PAR) of the intertidal zone (Graham 1996, Huovinen et al. 2000, Swanson & Druehl 2000),
44
A B C
45
D E F
Figure 3 Photographs of various Macrocystis populations. (A) Infrared aerial canopy photo of subtidal pyrifera-form population at La Jolla,
southern California. (Published with permission of Larry Deysher/Ocean Imaging.) (B) Shallow subtidal pyrifera-form population at Mar
Brava, central Chile. (Photograph by Michael Graham.) (C) Subtidal pyrifera-form population at Nightingale Island near Tristan da Cunha
Island, South Atlantic Ocean. (Published with permission of Juanita Brock.) (D) Intertidal integrifolia-form population at Van Damme State
Park, northern California. (Photograph by Michael Graham.) (E) Intertidal integrifolia-form population at Strait of Juan de Fuca, Washington.
(Photograph by Michael Graham.) (F) Intertidal integrifolia-form population at Huasco, northern Chile. (Photograph by Michael Graham.)
although wave activity, grazing and competition with other macroalgae in shallow subtidal areas
can also be important (Santelices & Ojeda 1984a, Foster & Schiel 1992, Graham 1997). At local
scales, decreased availability of light and rocky substratum, and occasionally sea urchin grazing,
appear to set the lower off-shore limits of Macrocystis populations (Pearse & Hines 1979, Lüning
1990, Spalding et al. 2003, Vega et al. 2005). Finally, within these upper and lower limits, the
lateral distribution of Macrocystis populations typically corresponds with abrupt changes in bathym-
etry or substratum composition (e.g., sand channels or harbour mouths; North & Hubbs 1968,
Dayton et al. 1992, Kinlan et al. 2005).
There is an interesting pattern within the global distribution of Macrocystis whereby different
regions may have large Macrocystis populations of one morphological form or another (Neushul
1971, Womersley 1987). For example, the integrifolia and angustifolia forms of Macrocystis are
generally found in shallow waters (low intertidal zone to 10 m depth), whereas the pyrifera form
is generally found in intermediate-to-deep waters (4–70 m depth) (Table 1). In the Northern
Hemisphere, the integrifolia form is most commonly observed at higher latitudes north of San
Francisco Bay with scattered populations found as far south as southern California (Abbott &
Hollenberg 1976, M.H. Graham, personal observations), whereas the pyrifera form is most common
at lower latitudes south of San Francisco Bay with scattered populations found as far north as
southeast Alaska (Gabrielson et al. 2000). In South America, the integrifolia and pyrifera forms
also appear to occupy shallow and deep habitats, respectively (Howe 1914, Neushul 1971). Lati-
tudinally, however, the Southern Hemisphere Macrocystis distribution is opposite that of the
Northern Hemisphere: the integrifolia form is generally found at lower latitudes, restricted to Perú
México, and northern Chile (Howe 1914, Neushul 1971), whereas the pyrifera form dominates the
higher latitudes of central and southern Chile (and Argentina; Barrales & Lobban 1975), but can
also be found far north in Perú (Howe 1914, Neushul 1971). The pyrifera form also appears to be
Table 1 Maximum depths of worldwide populations of Macrocystis ecotypes

Macrocystis form Location Depth (m) Reference
angustifolia South Australia 6 Womersley 1954
South Africa 8 Isaac 1937
integrifolia British Columbia 10 Druehl 1978
Northern Chile 8, 14 Neushul 1971, Vega et al. 2005
Perú 20 Juhl-Noodt 1958*
pyrifera Southern Chile 10 Dayton et al. 1973
Tasmania 15 Cribb 1954
New Zealand 16 Hay 1990
St. Paul/Amsterdam Is. 20 Delépine 1966*
Crozet Is. 25 Delépine 1966*
Falkland Is. 25 Powell 1981
South Georgia Is. 25 Skottsberg 1941
Southern California 30 Neushul & Haxo 1963
Central California 30 Spalding et al. 2003
Tristan da Cunha Is. 30 Baardseth 1941
Baja California 40 North 1971
Perú 40 Juhl-Noodt 1958*
Southern Argentina 55 Neushul 1971
Kerguelen Is. 40 Grua 1964*
Gough Is. 55 Chamberlain 1965
laevis Prince Edward Is. 68 Perissinotto & McQuaid 1992
* Depths interpreted by Perissinotto & McQuaid (1992).
46
most common where Macrocystis is found elsewhere in the Southern Hemisphere (e.g., Tasmania,
New Zealand, various sub-Antarctic islands), except in South Australia and South Africa where the
angustifolia form is common (Cribb 1954, Womersley 1954, 1987, Hay 1986, Stegenga et al. 1997).
Jackson’s (1987) analyses suggested that high latitude Macrocystis sporophytes would be light
limited in subtidal waters, forcing a shift in distribution to shallower water above 53° latitude. This
may explain the Northern Hemisphere distributional pattern, but cannot explain why shallow-water
Macrocystis is the most common form in northern Chile. Furthermore, exceptions to these patterns
clearly exist. For example, pyrifera-form individuals can be found in the intertidal zone (e.g., Guiler
1952, 1960 (Tasmania), Chamberlain 1965 (Gough Island), Westermeier and Möller 1990 (southern
Chile), van Tüssenbroek 1993 (Falkland Islands)), sometimes even side by side with integrifolia-
form individuals (M.H. Graham, personal observations in California; J.A. Vásquez, personal obser-
vations in northern Chile). Intermediate morphologies similar to the angustifolia form of South
Australia-South Africa can also be observed at intermediate depths (2–6 m) between adjacent
pyrifera-form and integrifolia-form populations in central California (M.H. Graham, personal
observations). Still, these global distribution patterns support the general consideration of the
integrifolia and angustifolia forms as having more shallow-water affinities than the pyrifera form.
Another interesting global distributional pattern is the apparent restriction of large Macrocystis
forests (>1 km2) to the southwest coast of North America (Point Conception in southern California
to Punta Eugenia in Baja California, Mexico; Hernández-Carmona et al. 1991, North et al. 1993),
although Macrocystis forests on most of the sub-Antarctic islands have not been explored. The
southwest coast of North America has broad shallow-sloping subtidal rocky platforms to support
wide Macrocystis populations (up to 1 km width), whereas the regions north to Alaska and south
to Patagonia have steep shores and typically support very narrow Macrocystis populations (<100 m
width); in some cases, narrow Macrocystis populations can fringe entire islands in the Pacific
Northwest (Scagel 1947), southern Chile (Santelices & Ojeda 1984b) and many sub-Antarctic
islands (e.g., Crandall 1915, Cribb 1954, van Tüssenbroek 1993). Thus, several key unanswered
questions remain: (1) does the geological restriction of Macrocystis to small forests outside southern
California affect the ecology of these systems (see ‘Population’ section, p. 54), (2) why are the
shallow-water forms found poleward in the Northern Hemisphere and equatorward in the Southern
Hemisphere, (3) does the recruitment of Macrocystis individuals to different depths or regions
determine their ultimate morphological form or (4) does variability in Macrocystis morphological
form determine the depth or region in which sporophyte recruitment and survival will be successful?
Life history
As with all kelps, Macrocystis exhibits a biphasic life cycle in which the generations alternate
(Sauvageau 1915), and the general life history is well understood (Figure 4; see review by North
(1994)). Macroscopic sporophytes attach to substrata by a holdfast consisting of a mass of branched
and tactile haptera. Dichotomously branched stipes arise from the holdfast and are topped by apical
meristems that split off laminae (blades) as they grow to the surface; gas-filled pneumatocysts join
laminae to the stipes and buoy them. The resulting fronds consisting of stipes, laminae and
pneumatocysts can form extensive surface canopies and represent the bulk of photosynthetic
biomass (North 1994). Other, shorter stipes give rise to profusely and dichotomously branched
specialised laminae near the base of the sporophyte (sporophylls) that bear sporangia aggregated
in sori (Neushul 1963); occasionally sori are observed on laminae in the canopy (A.H. Buschmann,
personal observations in southern Chile) and sporophylls can bear pneumatocysts (Neushul 1963).
Each sporangium contains 32 haploid biflagellate pyriform zoospores produced through meiosis
and subsequent mitoses (Fritsch 1945). Haplogenetic sex determination apparently results in a
1:1 male-to-female zoospore sex ratio (Fritsch 1945, Reed 1990, North 1994), although the two
47
Germlings
(drifting)
Female
gametophytes
Adults
Male (drifting)
gametophytes
Embronic
sporophytes
Zoospores
Adults
Juveniles (reproductive)
(recruits)
Adults
(attached)
Adults
(sterile)
Local population Remote populations
Figure 4 Macrocystis life cycle depicting various life-history stages important in regulating local Macrocystis
population dynamics. Ovals represent benthic stages and rectangles represent pelagic stages; white stages are
microscopic and shaded stages are macroscopic. Circular arrows represent potential for retention within
particular stages for unknown durations.
sexes cannot be distinguished easily at the zoospore stage (Druehl et al. 1989). Zoospores (~6–8
µm length) are released into the water column where they disperse via currents until they reach
suitable substrata where they settle, germinate and develop into microscopic male or female
gametophytes. As gametophytes mature, the females extrude oogonia (eggs) accompanied by the
pheromone lamoxirene (Maier et al. 1987, 2001). Upon sensing the pheromone, male gametophytes
release biflagellate non-photosynthetic antherozoids (sperm) that track the pheromone to the
extruded egg. Subsequent fertilisation gives rise to microscopic diploid sporophytes, which ulti-
mately grow to macroscopic (adult) size and complete the life cycle.
Although these steps necessary for Macrocystis to progress through its life cycle are straight-
forward, specific resources are necessary for gametogenesis, fertilisation, and growth of microscopic
stages. As a result, variability in environmental factors can greatly affect Macrocystis recruitment
success and completion of its life cycle. The experiments of Lüning & Neushul (1978) clearly
identified light quality and quantity as important in regulating female gametogenesis in Macrocystis,
and kelps in general. Deysher & Dean (1984, 1986a) quantified gross light (PAR), temperature and
nutrient (nitrate) requirements of Macrocystis gametogenesis and fertilisation, with embryonic
sporophyte formation limited to PAR above 0.4 µM photons (µEinsteins) m−2 s−1, temperatures
from 11 to 19°C and nitrate concentrations of >1 µM. Such critical irradiance, temperature and
nutrient thresholds were further supported by field experiments (Deysher & Dean 1986b). Although
these studies did not provide data amenable to the development of probability density functions
for predicting Macrocystis recruitment success as a function of variable environmental conditions,
the research was vital to the development of the concept of temporal ‘recruitment windows’, during
48
which environmental factors exceeded minimum levels for successful gametogenesis and fertilisa-
tion. Deysher & Dean (1986a) also found that the growth of embryonic sporophytes to macroscopic
size was inhibited at low PAR and nitrate concentrations, but these PAR levels were higher than
the threshold for gametogenesis and fertilisation. This suggests that the growth of embryonic
sporophytes to macroscopic size may be a stronger bottleneck in the Macrocystis life history than
gametogenesis and fertilisation. The photosynthesis studies of Fain & Murray (1982) similarly
identified differences in physiology between Macrocystis gametophytes and embryonic sporophytes.
The process leading to Macrocystis recruitment from gametophytes can thus be divided into
two functionally different stages: (1) sporophyte production (gametogenesis and fertilisation, with
relatively lower light requirements) and (2) growth of sporophytes to macroscopic size (with rela-
tively higher light requirements). It follows that the timing and success of Macrocystis recruitment
will depend on both the duration of each stage and whether such durations can be extended to
allow for delayed recruitment (Figure 4) similar to the concept of seed banks for terrestrial plants
(Hoffmann & Santelices 1991).
Laboratory and field studies indicate that the sporophyte production stage is relatively short
(1–2 months) and rigid in its duration, suggesting limited potential for delayed Macrocystis recruit-
ment via gametophytes. In California, female Macrocystis gametophytes appear to have an initial
competency period of 7–10 days prior to gametogenesis (North 1987) and lose fertility after ~30 days
(Deysher & Dean 1984, Kinlan et al. 2003), whereas in Chile, laboratory culture studies under
ample light and nutrient conditions suggest that Macrocystis female gametophytes may remain
fertile for up to 75 days (Muñoz et al. 2004). However, Macrocystis gametophytes can apparently
survive indefinitely under ‘unnatural’ artificial light-quality conditions (i.e., red light only; Lüning &
Neushul 1978). In California, unfertilised female gametophytes older than ~30 days have limited
potential for fertilisation (Deysher & Dean 1986b) and thus recruitment, which was supported by
the laboratory studies of Kinlan et al. (2003). As indicated by Kinlan et al. (2003), however, the
necessary studies have not been done to determine whether this lack of fertilisation success is due
to senescence of female gametophytes or of their male counterparts. Also, it has been demonstrated
that zoospore swimming ability is correlated with germination success (Amsler & Neushul 1990)
and a similar mechanism may affect fertilisation of oogonia by antherozoids. The demonstration
of a shorter life-span (period of fertility) for Macrocystis male gametophytes relative to females
would suggest the potential for cross-fertilisation among different zoospore settlements via peren-
nial females.
Another well-known aspect of sporophyte production is the minimum density of settled
zoospores necessary for recruitment. Specifically, the reliance of kelps on the presence of lamox-
irene as a trigger for antherozoid release (Maier et al. 1987, 2001) and the dilution of this sexual
pheromone over short distances from the oogonia inherently require sufficient zoospore settlement
densities (and survivorship to maturation) to ensure that males and females are close enough for
fertilisation to be successful. Such ‘critical settlement densities’ were demonstrated in a series of
laboratory and field experiments by Reed and his colleagues (Reed 1990, Reed et al. 1991).
Specifically, Reed et al. (1991) identified 1 settled zoospore mm−2 (vs. 0.1 or 10 settled zoospores
mm–2) as the minimum Macrocystis (and Pterygophora) zoospore settlement density above which
fertilisation and sporophyte production could be expected. These experiments focused on recruit-
ment from single zoospore settlement cohorts and cross-fertilisation among different zoospore
settlements may result in fertilisation even if cohort settlement densities are <1 settled zoospore
mm−2. It has recently been demonstrated that Macrocystis sporophytes can be produced from
unfertilised gametes through apogamy (Druehl et al. 2005). Although the frequency of partheno-
genic sporophyte production in the field has not been tested, parthenogenesis may obviate the need
for >1 settled zoospore to yield an adult sporophyte. Reed (1990) also demonstrated that species-
dependent female maturation rates combined with species-independent pheromone activity might
49
result in chemically mediated competition among microscopic stages of kelp species, although this
was only suggested for Macrocystis and Pterygophora in southern California. In addition to
producing valuable life-history data, these studies clearly demonstrated the utility of combining
laboratory and field experiments of kelp recruitment and resulted in a surge in studies of the ecology
of kelp microscopic stages.
Nevertheless, several key issues regarding the Macrocystis life history remain to be resolved.
Most importantly, Macrocystis microscopic life-history stages have not been observed in the field.
Microphotometric techniques have recently been developed for identifying Macrocystis zoospores
based on species-specific zoospore absorption spectra (Graham 1999, Graham & Mitchell 1999).
Subsequent determination of Macrocystis zoospore concentrations from in situ plankton samples
led to direct studies of Macrocystis zoospore planktonic processes (e.g., Graham 2003). However,
upon settlement, Macrocystis zoospores germinate into gametophytes of variable cell number and
pigment concentration, negating the use of microphotometric techniques for studying post-
settlement processes (Graham 2000). Fluorescently labelled monoclonal antibodies have been
developed for distinguishing between Macrocystis and Pterygophora gametophytes based on cell
surface antigens (Hempel et al. 1989, Eardley et al. 1990). However, the effectiveness of these tags
diminishes when applied to field samples, in which kelp cells are universally coated with bacteria
(D.C. Reed, personal communication). Additionally, although Kinlan et al. (2003) observed plas-
ticity in growth of laboratory-cultured Macrocystis embryonic sporophytes under realistic environ-
mental conditions (light and nutrients), and thus the potential for arrested development in this stage,
their experiments provided no evidence of arrested development of gametophytes. This study
demonstrated (1) that delayed recruitment of Macrocystis post-settlement stages is possible and
(2) the general lack of understanding of the physiological processes that regulate the growth,
maturation and senescence of Macrocystis microscopic stages. For example, it is considered that
kelp female gametophytes living under adequate environmental conditions will have only one or
very few cells, one oogonium per gametophyte, and become reproductive in the shortest period
possible (e.g., Lüning & Neushul 1978, Kain 1979). In the absence of light and nutrients, female
gametophytes are typically sterile and multicellular (e.g., Lüning & Neushul 1978, Kain 1979,
Hoffmann & Santelices 1982, Hoffmann et al. 1984, Avila et al. 1985, Reed et al. 1991), suggesting
a trade-off or antagonistic relationship between gametophyte growth and fertility. In some Chilean
populations, however, female Macrocystis gametophytes grown under standard laboratory condi-
tions (1) were multicellular, (2) produced multiple viable oogonia per gametophyte, (3) often
resulted in numerous sporophytes per gametophyte and (4) took longer to mature than Californian
populations (Muñoz et al. 2004). These results must be validated by additional laboratory and field
studies but they did demonstrate the highly plastic physiology of Macrocystis life-history stages.
Our lack of understanding of variability in the biology of Macrocystis microscopic stages, especially
at the global level, is an important constraint on future progress in Macrocystis population dynamics
(see ‘Population’ section, p. 54).
Growth, productivity and reproduction

Recruitment processes are the main determinant of when and where Macrocystis sporophytes might
occur, yet it is the survival and growth of established sporophytes that constrain sporophyte size,
self-thinning, population cycles and the primary productivity and canopy structure that ultimately
provide energy and habitat for Macrocystis communities.
The maximum age of Macrocystis sporophytes is unknown. Individual fronds generally senesce
after 6–8 months (North 1994) although van Tüssenbroek (1989c) observed maximum frond survival
of 1 yr and Macrocystis sporophytes can produce new fronds from apical meristems (frond initials)
retained above the holdfasts near the sporophylls (Lobban 1978a,b, van Tüssenbroek 1989c, North
50
1994). As such, the sporophytes may survive as long as they remain attached to the substratum
and environmental conditions are adequate for growth. In some regions of central California and
Argentina, most Macrocystis sporophytes die within a year due to high wave activity (Barrales &
Lobban 1975, Graham et al. 1997), whereas in southern California, sporophytes can live up to
4–7 yr (Rosenthal et al. 1974, Dayton et al. 1984, 1999), a life-span that coincides with the periodicity
of the El Niño Southern Oscillation (ENSO); in southern Chile, the life-span of Macrocystis
sporophytes often exceeds 2 yr (Santelices & Ojeda 1984b, Westermeier & Möller 1990). Interest-
ingly, the only Macrocystis populations known to recruit and senesce on an annual cycle occur in
the protected waters around 42°S in Chilean fjords (Buschmann et al. 2004a). The life-span of
vegetatively reproducing Macrocystis sporophytes (e.g., angustifolia and integrifolia forms) has
never been determined in the field, although integrifolia-form sporophytes have been shown to
survive very high levels of rhizome fragmentation (Druehl & Kemp 1982, Graham 1996) and
cultivated sporophytes can live 2–3 yr (Druehl & Wheeler 1986). In any case, the life-span of
Macrocystis sporophytes appears to be far less than that of other perennial kelp genera (Reed et al.
1996, Schiel & Foster 2006), for example, Pterygophora and Eisenia, which can live for 10+ yr
(Dayton et al. 1984).
The relatively high turnover of Macrocystis sporophytes is probably due to their massive size
(up to 400 fronds per pyrifera-form sporophyte; North 1994) and the almost strict reliance of
sporophyte growth and productivity on the biomass of the surface canopy (Reed 1987, North 1994,
Graham 2002). Shallow-water Macrocystis sporophytes typically have lower frond numbers than
deeper sporophytes (North 1994). Numerous studies have demonstrated high Macrocystis frond
productivity rates with estimates of 2–15 g fixed carbon m−2 day−1 in the Northern Hemisphere
(reviewed by North 1994), and values that vary between 7 and 11 g C m−2 day−1 in the southern
Indian Ocean (Attwood et al. 1991). Delille et al. (2000) also observed a significant ‘draw-down’
of pCO2 when off-shore water entered a dense Macrocystis bed at the Kerguelen Islands, suggesting
that the productivity of Macrocystis fronds was high enough to decrease inorganic carbon concen-
trations in the water column. Furthermore, Schmitz & Lobban (1976) determined that Macrocystis
sporophytes can translocate photosynthates from production sources in the surface canopy to energy
sinks (meristems, holdfasts, sporophylls) at rates of 55 to 570 mm h−1; the canopy typically
represents the greatest contribution to total sporophyte biomass (Nyman et al. 1993, North 1994).
Such high rates of productivity and translocation appear to be necessary to maintain sporophyte
growth in the face of high metabolic demands (Jackson 1987) because, unlike other perennial kelp
genera (e.g., Pterygophora), Macrocystis sporophytes have very limited nutrient and photosynthate
storage capabilities (2 wk; Gerard 1982, Brown et al. 1997). The subsequent reliance on the surface
canopy, and the vulnerability of surface canopy fronds to both physical and biological disturbance,
results in considerable spatial and temporal variability in Macrocystis productivity potential, size
structure and overall health.
The linkage between Macrocystis sporophyte growth, productivity and biomass therefore results
in a plastic response of sporophyte condition to temporal and spatial variability in resource avail-
ability (Kain 1982, Reed et al. 1996). The low storage capabilities are clearly disadvantageous
during periods of suboptimal environmental conditions, such as occur seasonally in southern
California (Zimmerman & Kremer 1986) and the inland waters of southern Chile (Buschmann
et al. 2004a). Again, other perennial kelp genera either possess greater storage capabilities or exhibit
seasonally offset periods of growth and photosynthesis in order to weather periods of low resource
availability (e.g., light or nutrients; Chapman & Craigie 1977, Gerard & Mann 1979, Dunton &
Jodwalis 1988, Dunton 1990). At high latitudes, like British Columbia, southeast Alaska, and the
Kerguelen and Falkland Islands, Macrocystis sporophyte growth follows distinctly seasonal patterns
in insolation, with frond elongation ranging from 2 to 4.7 cm day−1 during the summer maximum
(Lobban 1978b, Asensi et al. 1981, Druehl & Wheeler 1986, Wheeler & Druehl 1986, Jackson
51
1987, van Tüssenbroek 1989d). At lower latitudes, like California, distinct seasonal growth patterns
due to variability in insolation were not apparent (North 1971, Wheeler & North 1981, Jackson
1987, Gonzalez-Fragoso et al. 1991, Hernández-Carmona 1996). Instead, Zimmerman & Kremer
(1986) described seasonal frond growth rates that corresponded with variability in ambient nutrient
concentrations (nitrate), in which frond growth was maximised during winter-spring (12–14 cm
day−1; upwelling periods) and minimised during summer-fall (6–10 cm day−1; non-upwelling peri-
ods). In New Zealand, minimum Macrocystis frond growth rates also occurred during summer, but
were relatively high throughout the remainder of the year (Brown et al. 1997), whereas in northern
Chile frond growth rates of 5–10 cm day−1 were observed with no seasonal variability (Vega et al.
2005). In many regions, light and nutrients can be present well above limiting levels throughout
the year (e.g., central California or central Chile) thereby permitting continuously high Macrocystis
sporophyte productivity (Jackson 1987).
The reliance of Macrocystis sporophyte growth and productivity on the biomass and health of
the canopy also helps to explain much of the sensitivity of Macrocystis to ENSOs, relative to that
of other kelp genera (Dayton et al. 1999). There is a strong inverse relationship between water
column nitrate concentrations and water temperature (Zimmerman & Robertson 1985, Tegner et
al. 1996, 1997, Dayton et al. 1999, Hernández-Carmona et al. 2001). Kelp growth becomes nutrient
limited below approximately 1 µM nitrate, which typically occurs in southern California when
water temperatures rise above 16°C (Jackson 1977, Zimmerman & Robertson 1985, Dayton et al.
1999); the same threshold appears to occur around 18°C in Baja California, Mexico (Hernández-
Carmona et al. 2001). During ENSOs, depression of the thermocline shuts down nutrient replen-
ishment via coastal upwelling and decreases the propagation of nutrients via internal waves (Jackson
1977, Zimmerman & Robertson 1985, Tegner et al. 1996, 1997). Due to its limited nutrient storage
capabilities, Macrocystis canopy biomass begins to deteriorate when tissue nitrogen drops below
1.1% dry weight (Gerard 1982). When frond losses exceed frond initiation, the biomass necessary
to sustain meristems is lost and the sporophytes die. Sporophyte mortality was 100% in many
Macrocystis forests in southern and Baja California following the 1983 and 1997 ENSOs (Dayton
et al. 1984, 1992, 1999, Tegner & Dayton 1987, Dayton & Tegner 1989, Hernández-Carmona et al.
1991, Ladah et al. 1999, Edwards 2004), although sporophytes may find refuge in deep water
(Ladah & Zertuche-Gonzalez 2004) or within the benthic boundary layer (Schroeter et al. 1995).
Finally, during ENSOs, regulatory control over growth of juvenile Macrocystis sporophytes shifts
from light inhibition under Macrocystis surface canopies (Dean & Jacobsen 1984) to nutrient
limitation (Dean & Jacobsen 1986).
Extensive plasticity in sporophyte growth is by no means restricted to Macrocystis adults. Due
to the high temporal variability in sporophyte growth potential and the striking differences in
biomass among small and large Macrocystis sporophytes, the transition among different size classes
can also be delayed in time similar to the arrested development described above for embryonic
sporophytes. Santelices & Ojeda (1984a) and Graham et al. (1997) observed that Macrocystis
juveniles could survive for many months under adult canopies, growing rapidly to adult size when
adult densities decreased and light became available. Presumably, light levels under the canopy
were adequate to meet the metabolic demands of the juveniles, but inadequate to sustain growth
(Dean & Jacobsen 1984). It is unknown, however, how long juveniles or subadults can survive
such conditions. Another important feature of Macrocystis growth potential is that frond initiation
is indeterminate because sporophytes can tolerate sublethal biomass loss (loss of fronds) as long
as meristems are present and abiotic conditions are conducive to survival (North 1994). Subse-
quently, sporophyte age is decoupled from sporophyte size, which can be advantageous for both
young and old individuals, but disadvantageous to researchers trying to use size as a proxy of age
(Santelices & Ojeda 1984b). Graham (1997) found that large Macrocystis sporophytes living in
the surf zone suffered greater mortality due to wave action than those that survived sublethal loss
52
of canopy biomass, which presumably decreased overall sporophyte drag and the likelihood of
detachment by waves. Finally, it has been demonstrated that the response of Macrocystis juvenile
growth to variable nutrient concentrations is under genotypic control (Kopczak et al. 1991), resulting
in broad latitudinal variability in sporophyte growth and recruitment potential. Again, it is interesting
that such genotypic variation can occur in spite of non-trivial gene flow among Macrocystis
populations (Coyer et al. 2001).
The reliance of sporophyte growth on surface canopy biomass also constrains reproductive
output. Unlike most other kelp genera, Macrocystis sporophyll and sorus production can occur
continuously given adequate translocation of photosynthates from the surface canopy (Neushul
1963, McPeak 1981, Reed 1987, Dayton et al. 1999, Graham 2002, Buschmann et al. 2006). The
number of sporophylls per fertile Macrocystis sporophyte varies from 1 to 100+ (Lobban 1978a,
Reed 1987, Reed et al. 1997, Buschmann et al. 2004a, 2006), although sporophyll growth rates
have yet to be determined. In Macrocystis, two processes lead to turnover of reproductive material:
growth of sporophylls and production of sori on the sporophylls (Neushul 1963). Both processes
decrease in magnitude following either natural or experimental loss of canopy biomass (Reed 1987,
Graham 2002), although the cessation of sorus production appears to be more sensitive than
sporophyll growth to biomass loss and can result in complete sporophyte sterility within 9 days of
disturbance to the canopy (Graham 2002).
It is unknown whether sublethal biomass loss also affects the quantity or quality of zoospores
in sori or the timing of their ultimate release into the water column. Due to the continuous reliance
of Macrocystis reproduction on canopy biomass, however, variability in environmental factors can
also greatly affect reproductive output. Reed et al. (1996) demonstrated that nitrogen content of
Macrocystis zoospores varied as a function of in situ water temperature (and presumably water
column nutrient concentrations) and nitrogen content of adults, whereas the nitrogen content of
Pterygophora zoospores remained relatively constant. Reed et al. (1996) argued that the ability
of Macrocystis sporophytes to respond to favourable environmental conditions allowed them to be
reproductively successful despite their relatively short life-span. Again, such plasticity in reproduc-
tive timing can be adaptive, especially given the apparently low cost of reproduction in kelps
(DeWreede & Klinger 1988, Pfister 1992). For example, Macrocystis sporophytes living in wave-
exposed locations in southern Chile reproduce year-round and produce high numbers of sporophylls,
whereas Macrocystis sporophytes living in nearby wave-protected populations are annuals, have
increased zoospore production per sorus area and are fertile for only a few months, presumably to
ensure successful zoospore settlement and fertilisation prior to the disappearance of adult plants
every autumn (Buschmann et al. 2004a, 2006).
Overall, Macrocystis sporophyte growth, productivity and reproduction are very responsive to
variability in environmental conditions. This response differs from that of most known kelps and
other algae (see review by Santelices 1990) and is probably essential to the success of Macrocystis
as a competitive dominant throughout much of its global distribution. What remains to be deter-
mined, however, is how this variable physiology is expressed among the different morphological
forms of Macrocystis and across the variety of habitats in which Macrocystis populations are
present. For example, the integrifolia and pyrifera forms inhabit low intertidal and deeper subtidal
environments, respectively, which differ strikingly in factors known to regulate Macrocystis growth,
productivity and reproduction (e.g., water motion, water quality and light availability). Conse-
quently, it is expected that these two forms will respond differently to environmental perturbations
(e.g., van Tüssenbroek 1989c,e), with potentially significant consequences at the population and
community levels. This scenario is further complicated by the vegetative growth capabilities of the
integrifolia form, absent in the pyrifera form, because the relative contribution of vegetative growth
to sexual reproduction in maintaining integrifolia-form giant kelp populations is unknown. Further-
more, kelp physiological studies presently focus on measurements of physiological processes for
53
specific structures (e.g., photosynthesis, growth, or nutrient uptake of excised laminae), and few
have integrated these processes across entire sporophyte thalli (but see the translocation studies of
Schmitz & Lobban (1976)). For example, translocation elements (sieve tubes and trumpet hyphae)
run through the rhizomes of integrifolia-form sporophytes, which may spread over greater lengths
of substratum than pyrifera-form holdfasts, potentially providing a physiological connectivity
among fronds over the scale of metres. This limitation has inhibited the development of realistic
carbon and nitrogen budgets for kelps and thus constrained our understanding of the physiology
of entire sporophytes. This limitation is critical because it is at the level of individual sporophytes,
not individual laminae, that mortality, growth and reproduction have consequences for population
biology.
Population biology of Macrocystis

Most of the work on Macrocystis population dynamics prior to 1990 focused on processes regulating
seasonal-to-annual variability in adult sporophyte mortality (see review by North 1994), including
competition (Reed & Foster 1984, Santelices & Ojeda 1984a), herbivory (Harris et al. 1984,
Ebeling et al. 1985, Harrold & Reed 1985) and physical disturbance (Rosenthal et al. 1974, Dayton
et al. 1984). Stimulated by the research of Reed and his colleagues (Reed 1990, Reed et al. 1988,
1991, 1992, 1996, 1997, 2004), a more population-based approach to Macrocystis biology and
ecology has recently emerged in which studies have shifted to focus on reproduction, dispersal
and recruitment and the consequences of these processes to the persistence of Macrocystis popu-
lations. Subsequently researchers have developed a more integrated view of Macrocystis population
dynamics that unites variability in mortality agents with recruitment processes to provide a better
understanding of local and global differences in Macrocystis population cycles.
Stage- and size-specific mortality

Macrocystis populations do not exhibit unbounded growth (Dayton 1985a, Foster & Schiel 1985,
North 1994). Although Macrocystis populations are probably never at equilibrium, Macrocystis pop-
ulations often reach an apparent maximum in abundance or biomass per unit area (carrying capacity)
that is determined by the availability of environmental resources (e.g., space, light and nutrients;
Nisbet & Bence 1989, Burgman & Gerard 1990, Graham et al. 1997, Tegner et al. 1997). Further-
more, it has been well established that a variety of density-dependent and density-independent
processes result in stage- and size-specific sporophyte mortality (reviewed by Schiel & Foster 2006)
and retain Macrocystis at a population level below carrying capacity and initiate population cycling.
Due to their large size and high drag, Macrocystis adult sporophytes are extremely vulnerable
to removal by high water motion, and wave-induced sporophyte loss is considered the primary
factor resulting in Macrocystis mortality (Foster 1982, Dayton et al. 1984, Seymour et al. 1989,
Schiel et al. 1995, Graham 1997, Graham et al. 1997, Edwards 2004). The probability of a sporo-
phyte being removed from the substratum by passing or breaking waves increases when the force
(drag) experienced by the sporophyte due to water motion (related to both water velocity and the
cross-sectional area of the sporophyte exposed to the flow; Seymour et al. 1989, Utter & Denny
1996) exceeds the attachment strength of the sporophyte holdfast (for whole sporophyte mortalities)
or the breaking strength of individual fronds (for sublethal frond mortality; Utter & Denny 1996).
High seasonal and year-to-year variability in wave intensity and sporophyte biomass therefore
results in highly variable sporophyte mortality throughout the year. For example, in California,
most sporophyte mortalities occur during the first large fall-winter storms (Zobell 1971, Gerard
1976, Graham et al. 1997), when adult biomass is high following long periods of low wave activity
(spring to fall). It appears that sporophytes that survive these storms, but shed fronds and canopy
54
biomass, decrease their overall drag and increase the probability of surviving subsequent and often
more severe storms (Graham et al. 1997). On the Chatham Islands, Macrocystis populations are
only found at protected sites (Schiel et al. 1995) and never attain large sporophyte or population
sizes. In southern California, uprooted sporophytes are often observed entangled with attached
sporophytes, further increasing the attached sporophytes’ drag and probability of detachment
(Rosenthal et al. 1974, Dayton et al. 1984) and resulting in a ‘snowball effect’ that can clear large
swaths in the local population (Dayton et al. 1984). Such massive entanglements, however, appear to
be rare in central California (Graham 1997), possibly due to more rapid transport of detached sporo-
phytes out of and away from the local population. Increased sporophyte biomass, therefore, simul-
taneously increases both Macrocystis growth and reproductive potential (described in the Organ-
ismal biology section) and the probability of wave-induced mortality. This trade-off between fitness
and survival is probably viable because of the temporal and spatial unpredictability in wave intensity
experienced throughout the alga’s global distribution and its ability to survive and quickly recover
from sublethal loss of biomass. Exceptions are the wave-protected annual Macrocystis populations
in southern Chile in which there seems to be no trade-off between reproductive output and survival
(Buschmann et al. 2006). In this case, synchronous growth, reproduction and senescence occur in
the near absence of water motion.
Despite the high temporal variability in wave-induced mortality, Macrocystis sporophytes
exhibit distinct spatial patterns in survivorship. Wave-induced mortality of all size classes of adult
sporophytes increases with both increasing wave exposure (Foster & Schiel 1985, Graham et al.
1997) and decreasing depth (Seymour et al. 1989, van Tüssenbroek 1989c, Dayton et al. 1992,
Graham 1997). These patterns are primarily due to spatial variability in water motion because wave
activity increases toward shallow water, the tips of rocky headlands and regions of high storm
production (e.g., the relatively stable winter Aleutian low-pressure system in the Northern Hemi-
sphere). Graham et al. (1997), however, also observed that Macrocystis holdfast growth decreased
significantly along a gradient of increasing wave exposure, possibly due to greater disturbance to
the Macrocystis surface canopy, which reduces translocation to haptera and thereby reduces holdfast
growth (Barilotti et al. 1985, McCleneghan & Houk 1985). Thus, increased wave forces and
decreased strengths of holdfast attachment can act in combination to decrease Macrocystis sporo-
phyte survival; Graham et al. (1997) observed that Macrocystis sporophyte life-spans rarely
exceeded 1 yr at their most wave-exposed sites. Although all of these described patterns may
possibly exist for any Macrocystis life stage, the likelihood of wave-induced mortalities will be
much lower for the smaller life stages due to both decreasing thallus size and decreasing water
velocities within the benthic boundary layer. Additionally, other hydrographic factors can result in
high sporophyte mortalities in relatively wave-protected regions (e.g., tidal surge, nutrient limitation,
temperature and salinity stress; Buschmann et al. 2004a, 2006).
Biological processes also clearly play a role in mortality of Macrocystis sporophytes. During sea
urchin population outbreaks, sea urchin grazing of Macrocystis holdfasts can result in (1) detachment
of adult sporophytes and their removal from the population (Dayton 1985a, Tegner et al. 1995a),
(2) modification of sporophyte morphology (Vásquez & Buschmann 1997) and (3) removal of
entire recruits and juvenile sporophytes (Dean et al. 1984, 1988, Buschmann et al. 2004b, Vásquez
et al. 2006). Unlike some locations (e.g., the Aleutian Islands; Estes & Duggins 1995), widespread
destruction of Californian and Chilean Macrocystis populations by sea urchin grazing is rare
(Castilla & Moreno 1982, Foster & Schiel 1988, Steneck et al. 2002, Graham 2004). Still, sea
urchin outbreaks can result in episodic deforestation of Macrocystis populations up to a scale of a
few kilometres (Dayton 1985a). In healthy southern California systems, sea urchins can live in
Macrocystis holdfasts and result in holdfast cavitation and thus a decrease in sporophyte attachment
strength (Tegner et al. 1995a). Although episodic and small scale, the prevalence of holdfast cavitation
by sea urchins increases with increasing sporophyte age, thereby increasing the vulnerability of
55
large, older sporophytes to wave-induced mortality (Tegner et al. 1995a). Infestations of Macrocystis
sporophytes by epizoites and small herbivorous crustaceans (amphipods and isopods) have also
been observed worldwide (North & Schaeffer 1964, Dayton 1985b). Most outbreaks of herbivorous
crustaceans simply result in sublethal biomass loss (Graham 2002), which will effectively decrease
sporophyte drag and thus possibly wave-induced mortality. Crustacean infestations can also occur
in the holdfasts and result in increased mortality due to decreased sporophyte attachment strength
(North & Schaeffer 1964, Ojeda & Santelices 1984). When carnivorous ‘picker’ fishes are absent
from the water column in both California and Chile, outbreaks of epiphytic sessile invertebrates
(bryozoans, kelp Pecten spp., spirorbids) often result (Bernstein & Jung 1979, Dayton 1985b),
weighing down Macrocystis sporophyte canopies and either (1) increasing the likelihood of detach-
ment due to water motion or (2) bringing surface canopy biomass into contact with grazing activities
of benthic herbivores (Dayton 1985b). Although seemingly important, there are very few data
concerning the importance of these processes in regulating Macrocystis mortality worldwide.
Finally, although not a natural biological disturbance, human harvesting of Macrocystis canopies
does not appear to have significant effects on sporophyte survival (Kimura & Foster 1984, Barilotti
et al. 1985, Druehl & Breen 1986).
Inter- and intraspecific competition for space and light are important in regulating the survival
of Macrocystis microscopic stages (gametophytes and embryonic sporophytes) to macroscopic size
(juveniles; less than tens of centimetres), and growth of Macrocystis juveniles to adult size (Schiel
& Foster 2006). Smaller Macrocystis thalli are vulnerable to overgrowth by seaweeds and other
kelps (Santelices & Ojeda 1984a, Vega et al. 2005), and even by conspecifics in monospecific stands
(Schroeter et al. 1995, Graham et al. 1997). Intraspecific competition for space is likely to be most
severe at the smaller size classes because critical zoospore settlement densities will result in high
densities of microscopic embryonic sporophytes following fertilisation and the large size of adult
Macrocystis holdfasts (up to 1 m diameter) necessitates that many recruits and juveniles will be
smothered as nearby sporophytes grow in size. After Macrocystis sporophyte densities are initially
thinned by competition for space, competition for light increases as sporophytes begin to grow to
the water surface (Dean & Jacobsen 1984). Sporophytes that reach the surface will have enhanced
photosynthetic rates and be able to translocate more photosynthates to basal meristems for new
frond initiation (North 1994). As such, sporophytes that gain the competitive edge of a surface
canopy may become even larger, increasing their likelihood of outcompeting neighbours. Water
column nutrients further constrain the maximum amount of surface canopy biomass, apparently
regulating the total number of Macrocystis fronds per square meter (the frond carrying capacity;
North 1994, Tegner et al. 1997). The ontogenetic development of a Macrocystis cohort is, therefore,
dominated by self-thinning (Schiel & Foster 2006), in which high densities of small individuals
ultimately yield much lower densities of very large individuals (North 1994).
The applicability of this self-thinning model in Macrocystis populations, however, has not been
tested directly. North (1994) estimated the frond carrying capacity of a typical Macrocystis popu-
lation to be 10 fronds m−2, whereas Tegner et al. (1997) found frond carrying capacity to vary
according to oceanographic climate, being higher during cooler, nutrient-rich conditions (La Niña)
and lower during warmer, nutrient-poor conditions (El Niño). Schiel et al. (1995) also observed at
the Chatham Islands that a site with larger Macrocystis sporophytes had lower population densities
than a site dominated by smaller Macrocystis sporophytes. Many researchers have estimated that
self-thinning ultimately results in adult sporophyte densities of 1 per 10 m2 (Dayton et al. 1984,
1992, Graham et al. 1997), although the accuracy of this value has never been assessed experimen-
tally. Furthermore, these studies have been restricted to pyrifera form populations in central,
southern and Baja California. In other systems (e.g., Chile, New Zealand), pyrifera form individuals
do not grow to large sizes or form large populations (Schiel et al. 1995) and conspicuous self-
thinning of these populations has not been observed (Buschmann et al. 2004a, 2006). Similarly,
56
shallow-water integrifolia-form sporophytes exhibit vegetative propagation, resulting in coalescent

holdfasts, and the concept of sporophyte self-thinning may be irrelevant to these populations
(A. Vega & J.A. Vásquez, unpublished data).
As previously described, Macrocystis microscopic stages have high light requirements and are
thus highly vulnerable to inter- and intraspecific competition for light (Schiel & Foster 2006).
Due to their small size, Macrocystis gametophytes and embryonic sporophytes are also highly
vulnerable to sand scour (Dayton et al. 1984) and smothering by sediments (Devinny & Volse 1978)
and by macro- (Dean et al. 1989, Leonard 1994) and mesograzers (Sala & Graham 2002).
Finally, it should be noted that all of the above mortality agents typically result in small- to
mesoscale variability in the stage and size structure of Macrocystis populations. During normal
conditions, many factors are typically acting to regulate sporophyte survival in a probabilistic
fashion, resulting in high variability in sporophyte abundance and size structure at the scale of tens
to hundreds of metres (Edwards 2004). During episodic storms and ENSOs, however, multiple
factors (e.g., wave intensity and nutrient limitation) may act simultaneously to produce massive stage-
and size-dependent mortalities homogeneously over broad spatial scales of 10s to 100s of km
(Edwards 2004).
Dispersal, recruitment and population connectivity

The field ecology of microscopic life-history stages is perhaps the most dynamic and least under-
stood aspect of Macrocystis population biology (North 1994), and that of seaweeds in general
(Santelices 1990, Amsler et al. 1992, Norton 1992). Previous life-history studies for Macrocystis
indicate the potential for a wide variety of temporal and spatial variability in the time an individual
remains within a life-history stage, or the time necessary to proceed to subsequent stages (Figure 4).
This temporal flexibility in the life history begins with dispersal and ultimately results in variability
in recruitment and thus demographic interactions within a population (Santelices 1990).
Adult Macrocystis sporophytes typically produce zoospores with limited dispersal abilities (e.g.,
Anderson & North 1966, Dayton et al. 1984, Gaylord et al. 2002, Raimondi et al. 2004), suggesting
a tight coupling between zoospore output, dispersal and recruitment (Graham 2003). Recent studies,
however, have indicated that the supply of propagules of marine organisms can be decoupled from
the adult demographic and genetic patterns, as propagules are dispersed far from their natal site
(e.g., Roughgarden et al. 1988, Downes & Keough 1998, Wing et al. 1998, Shanks et al. 2000).
This decoupling also seems to apply to Macrocystis (Reed et al. 1988, 2004, 2006, Gaylord et al.
2002), especially when the populations are not large enough for modification of currents by the
canopy (Jackson & Winant 1983, Jackson 1998, Graham 2003). Because of their small size,
Macrocystis zoospores will clearly be transported as far as available currents advect them (Gaylord
et al. 2002). However, if adult sporophytes modify current directions and velocities, effective
zoospore dispersal can be decreased, coupling zoospore supply to relative changes in the density
and size structure of the adult sporophytes (Graham 2003). Subsequently, Macrocystis forests can
vary between ‘open’ and ‘closed’ populations, depending on their size, isolation and geographic
location (Graham 2003, Reed et al. 2004, 2006). Furthermore, Macrocystis zoospore dispersal can
be enhanced by episodic periods of high zoospore production that coincide with storms (Reed et al.
1988, 1997), large population sizes (and thus high source zoospore concentrations; Reed et al. 2004,
2006) and turbulent resuspension of zoospores within the benthic boundary layer (Gaylord et al.
2002). Together, spatial and temporal variability in water motion, zoospore output and Macrocystis
forest size results in high variability in the effective ranges of zoospore dispersal (Reed et al. 2006).
Nevertheless, it is likely that the dispersal dynamics described for a few large Macrocystis
forests in southern California are unique to this region (e.g., Point La Jolla and Point Loma at
1–8 km2; Dayton et al. 1984, Graham 2003, Reed et al. 2006) because most Macrocystis forests
57
worldwide are relatively small (<1 km2) and consist of narrow belts that fringe coastlines and
nearshore islands. In these cases, the retention of zoospores within the small natal adult populations
will be decreased, potentially reducing the probability of self-seeding of the populations and thus
increasing the reliance of the population on external propagule sources (Reed et al. 2004, 2006).
The potential for long-distance dispersal to effectively connect these small populations that occur
over broad regions (e.g., in central California, Chile, Australia, New Zealand) has not been tested
but models suggest that regional population connectivity via zoospore dispersal is likely (Reed
et al. 2006). Furthermore, alternative mechanisms for colonisation and population persistence
should be explored in these systems. For example, long-distance dispersal by means of drifting
sporophytes or reproductive fragments has been suggested as an important mechanism for Macro-
cystis colonisation (Figure 5; Anderson & North 1966, Dayton et al. 1984, Macaya et al. 2005,
Hernández-Carmona et al. 2006). Drifting reproductive sporophytes have been shown to be abun-
dant along broad regions of the Chilean and Californian coasts (Macaya et al. 2005, Thiel & Gutow
2005a, Hernández-Carmona et al. 2006), and drifting sporophytes can remain reproductively viable
in central California for over 125 days (Hernández-Carmona et al. 2006).
Clearly, dispersal distance alone cannot explain variability in local or remote recruitment,
including the colonisation of new substrata (Reed et al. 1988, 2004, 2006). Critical zoospore
settlement densities necessary for Macrocystis recruitment will inherently limit effective dispersal
distance to much less than the distance travelled by individual zoospores (Gaylord et al. 2002, Reed
et al. 2006). The key to long-distance colonisation, therefore, is not the arrival of a kelp propagule
to new substrata, but rather the arrival of two propagules (of opposite sex) within millimetres of
each other and their ultimate survival to sexual maturity. As such, new colonisations are rarely
Figure 5 (A) Drifting Macrocystis sporophyte, southern California. (Published with permission of Phillip
Colla/Oceanlight.com.) (B) Epi-fluorescent micrograph of drifting Macrocystis sporangia observed in water
sample (15 m depth) from Point Loma kelp forest, southern California (note individual zoospores with plastids).
(Photograph by Michael Graham.) Macrocystis identification based on species-specific spectrophotometric
signature (Graham 1999).
58
observed farther than tens of metres from individual Macrocystis sporophytes (Anderson & North
1966, Dayton et al. 1984, Reed et al. 2004, 2006) or hundreds of metres from Macrocystis popu-
lations (Anderson & North 1966, Reed et al. 2004, 2006). Physical and biological processes that
promote the arrival of zoospore aggregations to suitable substrata will, however, enhance the
frequency of long-distance colonisation. For example, Reed et al. (1997) observed a synchronous
decline in Macrocystis sorus area that was correlated with increased storm-induced water motion,
potentially indicating a synchronous bout of reproductive output. The locally increased density of
zoospores in the water column, and the high along-shore advection that occurs during such storms,
may help to extend the colonisation distance (Gaylord et al. 2002). Similarly, annual Macrocystis
populations in southern Chile exhibit increased zoospore production per soral area over short
reproductive periods, potentially increasing the temporal aggregation of settled zoospores (Busch-
mann et al. 2006). Other kelps also synchronise reproductive output (McConnico & Foster 2005),
increasing the likelihood that critical zoospore settlement densities will be exceeded, if only for a
short time. Drifting Macrocystis sporophytes may provide an additional aggregation mechanism
because reproductive sporophylls will travel together (Hernández-Carmona et al. 2006) and Dayton
et al. (1984) observed a path of recruitment in the trail of a drifting reproductive Macrocystis
sporophyte. Additionally, the detachment and dispersal of reproductive sporophylls, or even intact
sporangia (Figure 5), during periods of high reproductive output may also increase colonisation
distances as long as a high density of zoospores is released and they gain attachment to the
substratum. Benthic invertebrates that catch and eat such drifting reproductive fragments may
facilitate this process (Dayton 1985a).
In order to reach suitable settlement substratum, Macrocystis zoospores must enter the benthic
boundary layer where they respond to a chemically, physically and biologically heterogeneous
microenvironment (Amsler et al. 1992). At this microscale, zoospores can orient their movement
relative to nutrient gradients (Amsler & Neushul 1989) and settle preferentially in regions of high
micronutrient concentrations (Amsler & Neushul 1990); all kelp zoospores lack eyespots (Henry &
Cole 1982) and therefore are not phototactic (Müller et al. 1987). Energetic resources to support
zoospore swimming appear to come from a combination of zoospore photosynthesis and lipid
reserves (Reed et al. 1992, 1999). These experiments suggest an adaptation that enhances the
probabilities for settlement in suitable microenvironments for growth and reproduction of gameto-
phytes (Amsler et al. 1992). Upon settlement, the survival of Macrocystis gametophytes is low,
with <0.1% of the female gametophytes being fertilised (Deysher & Dean 1986a).
Microscopic stages, however, should not be considered simply an obstacle in the Macrocystis
life history that must be overcome in order for populations to persist. In fact, recent studies have
suggested that the microscopic stages may play a key role in population persistence by allowing
Macrocystis to survive environmental conditions that are unfavourable to macroscopic sporophytes.
Ladah et al. (1999) observed rapid and widespread Macrocystis recruitment following ENSO
1997–1998, which completely destroyed Macrocystis sporophytes over a 500-km region. The lack
of nearby reproductive adults and homogeneity in recruitment over this broad region suggested
that long-distance zoospore dispersal or individual drifting sporophytes were not the source of
recovery (although deep-water refuges were possible; Ladah & Zertuche-Gonzalez 2004). Ladah
et al. (1999) concluded that recruitment from persistent microscopic stages must have fuelled the
recovery, similar to the assumption by Buschmann (1992) that over-wintering microscopic stages
must link consecutive annual Macrocystis populations in southern Chile. Clearly, microscopic stages
of many kelp taxa can persist through adverse environmental conditions, although field studies by
Deysher & Dean (1986b) and Reed et al. (1997) have suggested that this is not true for Macrocystis.
Kinlan et al. (2003) recently demonstrated that the development of Macrocystis embryonic sporo-
phytes could be delayed under limited light and nutrients for at least 1 month. When resources
59
were restored, the surviving embryonic sporophytes grew quicker and reached larger sizes than
their ‘well-fed’ controls. Durations of arrested development >1 month were not explored, yet the
identification of Macrocystis embryonic sporophytes (rather than gametophytes) as a potentially
persistent stage may be important because high zoospore settlement densities are no longer neces-
sary for recruitment. In any case, the arrested development of Macrocystis microscopic stages
probably results from negligible growth due to inadequate resources (e.g., light or nutrients) rather
than a true physiological state of dormancy as in many terrestrial seed plants.
Numerous studies have identified low benthic irradiance as a key environmental factor limiting
Macrocystis recruitment (e.g., Dean & Jacobsen 1984, Reed & Foster 1984, Santelices & Ojeda
1984a, Deysher & Dean 1986b, Schroeter et al. 1995, Kinlan et al. 2003). In the field, such light
limitation along the deep limit of Macrocystis is typically due to poor water quality and high light
extinction with depth (e.g., Spalding et al. 2003). Between the shallow and deep limits, overlying
canopies of kelp, foliose and coralline algae regulate light available for Macrocystis recruitment
(e.g., Dean & Jacobsen 1984, Reed & Foster 1984, Santelices & Ojeda 1984a). Macrocystis
sporophytes that recruit to turf algae are typically removed by water motion before becoming firmly
attached to the substratum (Leonard 1994, Graham 1997). In fact, one of the few patterns to emerge
clearly for Macrocystis populations worldwide is that disturbances to Macrocystis canopies are
typically followed by Macrocystis recruitment (Dayton & Tegner 1984, Dayton et al. 1984, 1992,
1999, Reed & Foster 1984, Santelices & Ojeda 1984a, Graham et al. 1997). However, in annual
Macrocystis populations present in southern Chile, there is a time lag of 3–5 months between the
disappearance of the canopy and subsequent recruitment (Buschmann et al. 2006). This population
is also unique in that most Macrocystis sporophytes recruit to and grow upon the shells of large
filter-feeding slipper limpets (Crepidula; Buschmann 1992). Finally, Raimondi et al. (2004) have
recently demonstrated inbreeding depression (reduced growth) of Macrocystis recruits due to self-
seeding in close proximity to adult sporophytes. Thus, although most zoospores may only travel
short distances, inbreeding depression may select for cross-seeded recruits and enhance the effec-
tiveness of long-distance zoospore dispersal in driving within-population recruitment. The popula-
tion consequences of this intriguing result await exploration.
The vegetative propagation of integrifolia-form sporophytes following sporophyte recruitment
may enhance the persistence of Macrocystis populations, especially in the absence of consistent
zoospore supply. Buschmann et al. (2004a) observed low sporophyte fecundity in small and narrow
northern Chilean integrifolia-form populations relative to the larger central Chilean and pyrifera-
form populations, suggesting that sexual reproduction is less effective in these shallow-water
populations. Therefore, the role of dispersal in Macrocystis population dynamics must be considered
relative to the specific environmental and demographic contexts within which the populations exist,
especially with regard to population size and isolation, near-shore hydrodynamics, regeneration
capacity and differences in sexual and vegetative reproductive potential among Macrocystis forms.
It is important to note that recruitment to integrifolia-form populations is noticeably absent in
California (Setchell 1932, Graham 1996) but relatively common in British Columbia (Druehl &
Wheeler 1986) and northern Chile (Vega et al. 2005).
The ultimate consequence of 20+ yr of research on Macrocystis sporophyte mortality, propagule
dispersal and population recruitment has been the integration of available data to support the
functioning of regional Macrocystis forests as metapopulations (Reed et al. 2006). Reed et al. (2006)
estimated that the frequency of local extinctions and recolonisations for Macrocystis populations
in southern California occurred over broad temporal scales (months to 10+ yr). Local extinction
rates decreased with increasing population size and decreasing population isolation, with extinction
durations rarely exceeding 2 yr. Reed et al. (2006) also identified a broad spectrum of interpopu-
lation distances (hundreds of metres to tens of kilometres). It is suggested here that the broad array
of Macrocystis dispersal vectors, effects of local hydrodynamics, coupling of dispersal distances
60
to forest size and the potential persistence of microscopic life-history stages may again be advan-
tageous to maintaining demographic and genetic connectivity with Macrocystis metapopulations.
Demography and population cycles

Temporal variability in Macrocystis sporophyte abundance ranges from highly predictable to cha-
otic, depending on the spatiotemporal scales of interest. In some locations, sporophyte mortality
can been synchronised over broad spatial scales, typically driven by predictable seasonal mortalities,
such as in the wave-exposed regions of South America (Barrales & Lobban 1975, Dayton 1985b)
and California (Foster 1982, Dayton et al. 1984, Graham et al. 1997). Given the potential for
continuous Macrocystis recruitment, recruitment suppression by surface canopies, and the potential
for delayed recruitment, the response to these synchronised mortalities can be rapid and massive
(Dayton & Tegner 1984, Dayton et al. 1984) or delayed (Graham et al. 1997). Graham et al. (1997)
found that recruitment to adult size following winter mortalities was often delayed for many months
at wave-exposed sites on the Monterey Peninsula in California, due presumably to a lack of recruits
present to exploit the canopy opening. Although the cause of the recruitment delay was not
identified, Graham et al. (1997) suggested that production of new adults actually occurs in two
stages: first, the recruitment of macroscopic sporophytes (requiring both fertilisation and growth
to macroscopic size), and second, sporophyte growth to adult size. Nevertheless, recruitment appears
to drive Macrocystis population dynamics (Dayton et al. 1992, Graham et al. 1997, Buschmann
et al. 2006).
Graham et al. (1997) further suggested that the timing and magnitude of the disturbance
determined whether the recruitment stages occur in rapid succession or are separated by a delay.
For example, in the Point Loma kelp forest in southern California, Dayton et al. (1984, 1992, 1999)
have repeatedly observed massive recruitment to adult size following ENSOs (4- to 7-yr frequency).
In this case, pre-ENSO adult populations hover around carrying capacity with ENSOs typically
removing entire sporophytes from the majority of the population and allowing recruits to grow
quickly to adult size. In central California, however, the primary disturbance is caused by annual
storms that produce a mosaic of both lethal and sublethal mortalities (Graham et al. 1997, Edwards
2004), and the populations never truly reach carrying capacity. Individual sporophytes may be lost,
but canopies often recover quickly, decreasing the likelihood that recruits can grow directly to adult
size. As such, the massive synchronised recruitment that drives long-term cycles in Macrocystis
population dynamics in southern California (Dayton et al. 1984, 1992, 1999) may be typical of
regions that experience large, yet episodic disturbances (e.g., ENSO in California and Chile;
Edwards 2004, Vega et al. 2005), whereas regions that experience more chronic annual disturbances
may experience more unpredictable population cycling, such as in California (Graham et al. 1997)
and Chile (Buschmann et al. 2004a). Such a generalisation is consistent with the population
modelling studies of Nisbet & Bence (1989) and Burgman & Gerard (1990).
The effect of ENSO on Macrocystis population cycling also appears to vary among ENSOs.
Both ENSO-induced storms and nutrient deprivation are major sources of Macrocystis mortality
(Dayton et al. 1984, 1992, 1999, North 1994, Edwards 2004). During the 1982–1983 ENSO in
California, large storms preceded the period of anomalously warm temperature and high nutrient
stress (Dayton & Tegner 1984, Dayton et al. 1984), decimating Macrocystis populations throughout
their range (Tegner & Dayton 1987). During the 1997–1998 ENSO, however, southern and Baja
California kelp populations deteriorated in anomalously warm temperature prior to the massive
winter storms of 1998 (Edwards 2004). As a result, some sporophytes survived in deeper water
(Dayton et al. 1999, Edwards 2004, Ladah & Zertuche-Gonzalez 2004), potentially due to the
decreased drag of sublethal loss of canopy biomass. Again, in both cases, the combination of open
space cleared by storms, reduced canopy shading and subsequent La Niña conditions led to intense
61
recruitment in the spring (Dayton et al. 1984, 1992, 1999). Increased abundance of understory kelps
(e.g., Pterygophora californica, Laminaria farlowii and Eisenia arborea), usually inferior compet-
itors to Macrocystis, became well established during ENSO, persisted for many years post-ENSO
(Dayton & Tegner 1984, Dayton et al. 1984, 1999) and was shown to suppress Macrocystis
recruitment in local areas (e.g., Edwards & Hernández-Carmona 2005).
In the southeast Pacific, Soto (1985) also reported a massive mortality of Macrocystis from
18 to 30°S during ENSO 1982–1983, resulting in a collapse of the kelp harvest from 1983 to 1986
in northern Chile (National Fishery Service, SERNAPESCA 1980–1990). No such mortalities were
witnessed, however, in northern Chile Macrocystis populations during ENSO 1997–1998 (Vega
et al. 2005) where Macrocystis abundances were reduced but soon replaced by high recruitment.
The absence of an ENSO-induced Macrocystis collapse in northern Chile suggested (1) differential
effects of various ENSOs at different localities along the coastline; (2) presence of ‘source’ localities
(Camus 1994), which, due to certain attributes of the habitat, were able to maintain Macrocystis
populations that provided reproductive propagules to disturbed populations (‘sink’ localities); (3) the
existence of persistent microscopic life-history stages (Santelices et al. 1995) and (4) differential
effects of ENSO on intertidal versus subtidal Macrocystis populations. Unpredictably, Macrocystis
populations in northern Chile began to decrease following ENSO 1997–1998, apparently as a result
of La Niña 1999 (Vega et al. 2005). The direct cause remains unknown but is linked to Macrocystis
recruitment failure.
Ecology of Macrocystis communities

One of the most interesting aspects of any Macrocystis-dominated system is the linkage between
the dynamics and productivity of Macrocystis populations and the diversity and structure of their
associated floral and faunal communities. Indeed, the functional importance of Macrocystis within
giant kelp communities was apparent to even the earliest kelp forest ecologists (see, e.g., Darwin
1839). Here, recent advances in Macrocystis community ecology are explored through a discussion
of the structural role of Macrocystis within the system, resulting predator-prey interactions and
food web dynamics, and the effects of exploitation and global climate changes on the biodiversity
and stability of these coastal systems on global scales. The focus is on a mechanistic understanding
of Macrocystis systems; a more comprehensive treatment of Macrocystis community ecology can
be found in Foster & Schiel (1985).
Macrocystis as a foundation species

Macrocystis is the tallest benthic organism (Steneck et al. 2002). Due to their complex morphology,
Macrocystis sporophytes can alter abiotic and biotic conditions by dampening water motion (Jackson
& Winant 1983, Jackson 1998), altering sedimentation (North 1971), shading the sea floor (Reed &
Foster 1984, Edwards 1998, Dayton et al. 1999, Clark et al. 2004), scrubbing nutrients from the
water column (Jackson 1977, 1998), stabilising substrata (Neushul 1971, North 1971), providing
physical habitat for organisms both above and below the benthic boundary layer (reviewed by
Foster & Schiel 1985) and providing fixed carbon (from drift kelp to particulate and dissolved
organic carbon) within Macrocystis forests (Gerard 1976) and to surrounding habitats (reviewed
by Graham et al. 2003). The irony for Macrocystis community ecologists is that this complex role
of Macrocystis as the foundation of its associated community is both the impetus for mechanistic
community ecology studies and yet is the primary impediment to such studies.
There are three primary components to direct provision of habitat by Macrocystis sporophytes:
the holdfast, the mid-water fronds, and the surface canopy (Foster & Schiel 1985). Macrocystis
holdfasts are complex structures comprising numerous dichotomously branched and intertwined
62
haptera and are colonised by a highly diverse assemblage of algae, invertebrates and fishes (Figure
2A,B; Fosberg 1929, Andrews 1945, Cribb 1954, Ghelardi 1971, Jones 1971, Beckley & Branch
1992, Vásquez 1993, Thiel & Vásquez 2000). Haptera typically initiate from the primary stipe
dichotomy, with new haptera forming above older ones. The haptera generally grow until they
reach the substratum, thereby forming holdfasts that are initially 2-dimensional structures, and
depending on age, may ultimately become large 3-dimensional mounds. As the holdfasts grow,
new biomass accumulates along the outer surface, whereas the older biomass in the centre of the
holdfast becomes necrotic and cavitates (Cribb 1954, Ghelardi 1971, Tegner et al. 1995a). As such,
large holdfasts can provide different quantities and qualities of available habitat than smaller ones;
large holdfasts are generally restricted to angustifolia-, laevis- and pyrifera-form sporophytes
(Figure 2A), whereas the flat strap-like rhizomes of integrifolia-form sporophytes offer little habitat
to kelp forest organisms (Figure 2B; Scagel 1947).
Most work on Macrocystis holdfast communities has focused simply on species enumeration
(Ghelardi 1971, Jones 1971, Beckley & Branch 1992, Vásquez et al. 2001) and patterns of faunal
abundance and diversity as a function of holdfast size (Andrews 1945, Thiel & Vásquez 2000) or
time since dislodgement of holdfasts from the substratum (Vásquez 1993). Large holdfasts are
often encrusted with bryozoans and sponges and serve as refuges for crustaceans (e.g., amphipods),
molluscs, brittlestars and sea urchins, especially in the large cavitated centres of older holdfasts;
small holdfasts typically house the more mobile invertebrates (e.g., amphipods). Occasional her-
bivore outbreaks within Macrocystis holdfasts may contribute to sporophyte mortalities, especially
for large sporophytes (Jones 1971, Tegner et al. 1995a). Due to the dynamic nature of Macrocystis
populations, high variability in sporophyte size and intersporophyte distances may be of primary
importance in driving the abundance and diversity of holdfast communities within a population, as
predicted by ‘island biogeography’ theory (Thiel & Vásquez 2000). Nevertheless, it has not been
determined whether Macrocystis holdfast communities are of functional importance within the
larger kelp forest system.
The mid-water fronds and surface canopies are also host to a variety of fishes, sessile and
mobile invertebrates, and even birds and pinnipeds (reviewed by Graham 2004, Graham et al. 2007).
Encrusting bryozoans, hydroids and occasionally bivalves (Pecten) may cover large portions of
mid-water fronds (Scagel 1947, Wing & Clendenning 1971, Bernstein & Jung 1979, Dixon et al.
1981, Dayton 1985a,b, Hurd et al. 1994), which are inherently older than their surface-water
counterparts. The bulk of the faunal biomass in the mid-water, however, is locked up in crustaceans,
grazing molluscs (e.g., top and turban snails; Watanabe 1984a,b, Coyer 1985, 1987, Stebbins 1986)
and juvenile and adult fishes, which use the habitat as refuge, for foraging or as a focus of
aggregations (Bray & Ebeling 1975, Moreno & Jara 1984, Ebeling & Laur 1985, Hallacher &
Roberts 1985, DeMartini & Roberts 1990, Holbrook et al. 1990, Stephens et al. 2006). The
Macrocystis-fish association may be weaker, however, in areas with high relief (Stephens et al.
1984). Again, the importance of these faunal components to the system as a whole has not been
addressed.
The functional importance of Macrocystis canopies to the dynamics of the kelp forest commu-
nity, however, is well established. Macrocystis canopies are important recruitment sites for many
species of near-shore fishes (Carr 1989, 1991, 1994, Anderson 1994, Stephens et al. 2006), and the
direct link between canopy biomass, frond density or sporophyte density and fish abundance has
been demonstrated (Carr 1989, 1991, 1994, DeMartini & Roberts 1990, Holbrook et al. 1990,
Anderson 1994, 2001). These fish assemblages are important in controlling canopy herbivore
outbreaks (Bray & Ebeling 1975, Bernstein & Jung 1979, Dayton 1985a,b, Tegner & Dayton 1987,
Graham 2002), except in South America where canopy ‘picker-fish’ assemblages are apparently
absent (Dayton 1985b, Vásquez et al. 2006). Nevertheless, the Macrocystis-fish relationship is
complex. Some kelp forest fish taxa show a negative relationship with Macrocystis abundance
63
(e.g., Embiotoca lateralis), apparently due to the negative effect of Macrocystis on subsurface algal
assemblages that are important fish foraging habitats (Ebeling & Laur 1985, Holbrook et al. 1990).
It remains to be seen, however, whether such conspicuous habitat associations are specific to the
relatively benign regions in which they have been tested (e.g., southern California) or may also be
general to regions with higher spatial and temporal variability in Macrocystis population dynamics.
Furthermore, there appear to be no studies of fish-Macrocystis associations that have focused on
the shallow-water angustifolia or integrifolia forms.
Macrocystis sporophytes continue to provide habitat resources after detachment from the
substratum. Holdfast, mid-water fronds and canopies can retain epifaunal fishes and mobile and
sessile invertebrates even when drifting sporophytes travel long distances (Edgar 1987, Vásquez
1993, Helmuth et al. 1994, Hobday 2000a,b,c, Smith 2002, Macaya et al. 2005, Thiel & Gutow
2005a,b). In some cases, new species are added from the plankton to the communities on the
Macrocystis drifters (Hobday 2000c, Thiel & Gutow 2005b). Eventually the drifters (1) are depos-
ited onto coastal beaches where they can be buried or the pneumatocysts may break causing
sporophytes that are resuspended by high tides to be transported to deep water habitats (Zobell
1971), (2) lose their buoyancy at sea and sink directly to deep-water habitats (Harrold & Lisin
1989), (3) aggregate at convergence zones where they may grow vegetatively for undetermined
durations (North 1971, van Tüssenbroek 1989b, Thiel & Gutow 2005a) or (4) reconnect with coastal
Macrocystis populations (Hobday 2000a, Thiel & Gutow 2005b).
Additional habitat and trophic associations become apparent at the scale of Macrocystis pop-
ulations rather than individuals, probably due to the variety of ways that kelp forests affect the
near-shore environment. Kelp forests tend to be darker, less hydrodynamic habitats than adjacent
rocky reefs (Dayton 1985a, Foster & Schiel 1985) and it has been hypothesised that large kelp
forests may be sites where fixed carbon accumulates in the form of detritus (from drift kelp to
small particulates) (Gerard 1976, Harrold & Reed 1985, Graham et al. 2007). It is therefore not
surprising that up to 35% of 275 common kelp forest taxa (flora and fauna) in the southern California
Channel Islands were found to be associated with the presence of Macrocystis (Graham 2004),
25% of which were obligate associates; >90% of the taxa were more common in forested areas
than deforested areas. Many of these associates had either clear trophic linkages with Macrocystis
(e.g., abalones) or the associations were driven by habitat provision (e.g., kelp surfperch Brachy-
istius frenatus).
Similar forestwide associations are found for Macrocystis populations in northern Chile,
although species richness is much less than in California (Figure 6). Interestingly, the Chilean data
show first that the presence of other kelp taxa (e.g., Lessonia) can also drive changes in kelp forest
assemblage structure, and second that different species of kelp (e.g., Macrocystis vs. Lessonia) may
differ in the quality and quantity of habitat that they provision. Furthermore, Vega et al. (2005)
recently demonstrated that the morphology of the understory subtidal kelp Lessonia trabeculata
varies in the presence/absence of Macrocystis, potentially resulting in additional effects of Macro-
cystis distribution on community structure. Still, these studies have relied on natural kelp defores-
tations, for example due to sea urchin overgrazing, which can produce various factors that confound
variability in kelp presence. As such, the direct isolation of the importance of Macrocystis energy
and habitat provision relative to that of other kelps or non-kelp macroalgae species remains elusive
(Graham et al. 2007). Finally, due to the different growth rates and distribution of canopy biomass
among the different forms of Macrocystis (e.g., pyrifera vs. integrifolia forms), shallow- and deep-
water Macrocystis populations may also provide trophic and habitat resources in different ways.
For example, interfrond distances among shallow integrifolia- and angustifolia-form sporophytes
(Figure 2D) are much more homogeneous than their deeper pyrifera-form counterparts (Figure
2C), where high stipe densities are aggregated around individual sporophytes that are widely spaced.
Such ecotypic variability in the spatial distribution of suitable habitat (i.e., canopy fronds) may
64
affect the nature and strength of species associations in shallow versus deep water, especially among
fish taxa that preferentially use the canopy, mid-water fronds or water column spaces.
Trophic interactions and food webs

During his initial observations of Macrocystis forests in southern Chile, Darwin (1839) was struck
by the high diversity of species that appeared to be trophically linked to energy provision by
Macrocystis. He professed “The number of living creatures of all orders, whose existence intimately
depends on the kelp, is wonderful”. Indeed, all described Macrocystis forests harbour tens to
hundreds of species, most of which feed either on Macrocystis-derived fixed carbon (from direct
grazing to filter-feeding on Macrocystis detritus) or within some predatory trophic subweb founded
upon Macrocystis-based herbivores (Graham et al. 2007). Although food web studies are rare
(reviewed by Graham et al. in press), most species lists from within Macrocystis forests contain
taxa from multiple trophic levels, often with numerous taxa within each level, and typically a wide
variety of generalist and specialist species. It therefore seems unnecessary to review here the nature
of trophic interactions within Macrocystis forests. Instead, the focus is specifically on recent studies
of the fate of Macrocystis-based primary productivity and the importance of trophic interactions
in the dynamics and stability of Macrocystis forests because these are topics of global interest.
Gerard (1976) and Pearse & Hines (1976) estimated that although central Californian Macro-
cystis sporophytes had very high standing stock, most Macrocystis-based productivity entered the
food web through detrital pathways. Storm waves rip entire sporophytes from the substratum, break
fronds and erode senescent blades, sending detrital material of a wide size range to either the kelp
forest floor or out of the forest to other systems. Large detrital pieces (i.e., drift) collect near the
bottom where they are heavily grazed by asteroids, crustaceans (crabs, amphipods, isopods), snails
and fishes (Leighton 1966, Feder et al. 1974, Gerard 1976, Pearse & Hines 1976, Beckley & Branch
1992, Kenner 1992, Hobson & Chess 2001; see also reviews by Castilla 1985, Foster & Schiel
1985, Graham et al. 2007). When present, large pieces of Macrocystis drift make up the primary
diet of sea urchins (Leighton 1966, Castilla & Moreno 1982, Dayton et al. 1984, Vásquez et al.
1984, Castilla 1985, Ebeling et al. 1985, Harrold & Reed 1985). In Chile, Tetrapygus niger and
Loxechinus albus can catch drift in Macrocystis forests (Castilla 1985, Rodriguez 2003) but to a
lesser extent than Strongylocentrotus franciscanus or S. purpuratus in the northeast Pacific (Harrold
& Reed 1985, Harrold & Pearse 1987). When deprived of drift, S. franciscanus or S. purpuratus
abandon their normal ‘sit and catch drift’ strategy in search of attached algae (Mattison et al. 1977,
Ebeling et al. 1985, Harrold & Reed 1985, Harrold & Pearse 1987), a behavioural switch not
observed for other sea urchin taxa (Vásquez & Buschmann 1997, Steneck et al. 2002). Macrocystis
drift is also the main component of the diet of abalone in California (Leighton 1966, Tutschulte &
Connell 1988) and sea urchins and abalone are thought to compete strongly when Macrocystis drift
is in short supply (Tegner & Levin 1982). In the absence of drift, abalones often decrease in
abundance or disappear from the local system entirely (e.g., Graham 2004). Smaller detrital pieces
(i.e., particulate organ carbon, POC) make Macrocystis-based productivity accessible to many more
taxa (e.g., polychaetes, bivalves, sponges, crustaceans, ophiuroids, mysids or basically any kelp
forest detrital or filter feeder; Clarke 1971, Foster & Schiel 1985, Beckley & Branch 1992, Kim
1992, Graham 2004, Graham et al. 2007). Therefore, it is not surprising that sessile filter feeders
and mobile herbivores can be extremely diverse in Macrocystis forests (100+ taxa), with many taxa
disappearing during local Macrocystis deforestation (Graham 2004).
The direct grazing pathway is also utilised by a high diversity of kelp forest herbivores
(Rosenthal et al. 1974, Gerard 1976, Pearse & Hines 1976, Moreno & Sutherland 1982, Castilla
1985, Foster & Schiel 1985, Graham 2004, Graham et al. 2007). Snails, crustaceans, asteroids and
fishes can graze the benthic and water column biomass of Macrocystis sporophytes. These herbivores
65
Per cent cover
0 20 40 60 80 0 20 40 60 80 0 20 40 60 80 100
Mesophyllum sp.
Hildenbrandia sp.
Calcareous crust
Griffithsia chilensis
Glossophora kunthii
Green crust
Asparagopsis armata A
Gelidiales
Halopteris paniculata
66
0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 50
Porifera
Phragmatopoma
Spionidae
Balanus laevis
Austromegabalanus sp.
Bugula sp. B
Pyura chilensis
Vermetidae
Figure 6 Macroscopic species abundances in areas of sea urchin overgrazing (deforested), Lessonia trabeculata forests and Macrocystis (integrifolia-
form) forests for northern Chile (3–15 m depth). (A) Per cent cover of macroalgae; (B) per cent cover of sessile invertebrates.
Density (no. m−2)
0 10 20 30 40 50 0 10 20 30 40 50 0 10 20 30 40 50 60
Tetrapygus niger
Tegula atra
Tegula tridentata
Allopetrolisthes sp.
Nassarius gayi
Crassilabrum crassilabrum
Calyptraea trochiformis
Mitrella unisfasciata
Xantochorus cassidiformis
Prisogaster niger
Scurria sp.
Pagurus sp.
Ophiaycthis kroyerii C
Taliepus sp.
Tricolia mcleani
Abundance (no. 20 min transect−1)

0 2 4 6 8 0 2 4 6 8 0 2 4 6 8 10
Aplodactylus punctatus
67
Scarthichthys viridis
Doydixodon laevifrons
Cheilodactylus variegatus
Helcogrammoides cunninghami
Pinguipes chilensis
Hypsoblennius sordidus
Graus nigra
Chromis chrusma
Paralabrax humeralis
Trachurus murphyi
Isacia conceptionis
Prolatilus jugularis
Auchenionchus microcirrhis
Hemilutjanus macropthalmos
Mugil cephalus
Anisotremus scapularis
D
Calliclinus genicuttatus
Austromenidia laticlavia
Macrocystis Lessonia Deforested
Figure 6 (continued) (C) density of mobile invertebrates (no. m−2); (D) abundance (no. 20 min−1 of visual survey) of fishes. Data are means ± SD;
sample sizes range from 34 to 136. Data are from Nuñez & Vásquez (1987), Vásquez et al. (1998), Salinas (2000) and W. Stotz, J. Aburto &
L. Cailleaux, unpublished data.

generally have low per capita consumption rates (e.g., Jones 1971, Sala & Graham 2002) and
probably have little impact on Macrocystis standing stock, except during population explosions
(Jones 1971, Tegner & Dayton 1987, Graham 2002) or when Macrocystis sporophytes are small
in size (e.g., during recruitment or recovery following disturbance; Moreno & Sutherland 1982,
Harris et al. 1984, Castilla 1985, Sala & Graham 2002, Buschmann et al. 2004b). It is unknown
whether the diversity of these herbivores within Macrocystis forests may enhance or lessen the
effects of herbivory on Macrocystis survival and population dynamics, through complementarity
or competition, respectively.
It is well known, however, that sea urchins can have a great impact on Macrocystis standing
stock through direct grazing (Lawrence 1975, Pearse & Hines 1979, Schiel & Foster 1986, Harrold
& Pearse 1987, Vásquez & Buschmann 1997, Steneck et al. 2002, Vásquez et al. 2006). In some
systems (e.g., southern Chile), Macrocystis fronds can be weighted down by epizoites and sea
urchins can heavily graze water column biomass directly (Dayton 1985b). In most cases, however,
the greatest impact of sea urchin grazing on Macrocystis biomass is when sea urchins aggregate
on holdfasts and detach entire sporophytes, which then drift out of the system (North 1971, Foster
& Schiel 1985); sea urchins (and potentially other herbivores) may then keep the system in a
deforested state by grazing directly on Macrocystis recruits. Again, such overgrazing by Strongy-
locentrotus is apparently limited to periods of low drift availability (Ebeling et al. 1985, Harrold
& Reed 1985), which occur episodically at local scales within southern California (Foster & Schiel
1988, Steneck et al. 2002, Graham 2004). Sea otters (Enhydra lutris) inhibit sea urchin overgrazing
throughout the otter’s range (McLean 1962, Harrold & Pearse 1987, Foster & Schiel 1988, Watanabe
& Harrold 1991), which at present is limited mostly to the Northern Hemisphere north of Point
Conception (Laidre et al. 2001). Cowen et al. (1982) also observed that high wave action in central
California curtailed sea urchin foraging and allowed algal recovery. In this region, seasonal vari-
ability in wave intensity was suggested as the most important factor regulating the abundance and
structure of macroalgal assemblages (Cowen et al. 1982, Foster 1982).
The mechanisms controlling sea urchin overgrazing in southern California (south of Point
Conception), however, are controversial. In the absence of sea otters, various forms of abiotic and
biotic regulation of sea urchin populations have been proposed (see Foster & Schiel 1988, Steneck
et al. 2002). For example, storms and/or disease can wipe out large sea urchin aggregations over
relatively broad spatial scales (Ebeling et al. 1985, Tegner & Dayton 1991, Lafferty 2004), whereas
recruitment failure can limit replenishment of local sea urchin populations (Pearse & Hines 1987).
Nevertheless, the most popular explanation for the lack of large-scale sea urchin barrens in the
absence of sea otters in southern California is that other predators are controlling sea urchin abundance.
Various kelp forest predators eat sea urchins (see review by Graham et al. 2007), although most
only eat small sea urchins that are incapable of inflicting significant damage to Macrocystis
holdfasts. Sheephead (Semicossyphus), lobsters (Panulirus) and the sunflower stars (Pycnopodia)
appear to be the only Californian kelp forest predators other than sea otters that can feed on adult
sea urchins. In some southern California kelp forests, data suggest that sheephead and lobster
predation are important in controlling urchin abundance (e.g., Cowen 1983, Lafferty 2004) although
the diets of these species are highly variable in space and may not include sea urchins even when
sea urchins are present (e.g., Cowen 1986). Furthermore, in most cases the predation hypothesis
is invoked in the absence of field experimentation (but see Cowen 1983), which is problematic
since sheephead and lobsters have become relatively rare in southern California kelp forests (Dayton
et al. 1998), yet deforested areas are also relatively rare (Foster & Schiel 1988). Sea urchin
populations in southern California, therefore, are clearly regulated by multiple abiotic and biotic
processes, probably resulting in the low frequency of sea urchin barrens in the Southern California
Bight.
68
The only other Macrocystis systems for which sea urchin overgrazing has been observed are
in South America, although like California, large-scale overgrazing is rare (Vásquez & Buschmann
1997, Steneck et al. 2002). In northern Chile, large-scale overgrazing of Macrocystis by Tetrapygus
niger appears to be limited by high water motion in the region where Tetrapygus and Macrocystis
distributions overlap (Vásquez & Buschmann 1997). Sea urchin overgrazing in northern Chile is
subsequently restricted to particular depth zones. Also, the asteroids Luidia and Meyenaster are
solitary hunters within subtidal habitats and are important predators on Tetrapygus niger and other
echinoids and asteroids (Viviani 1979, Vásquez & Buschmann 1997, Vásquez et al. 2006). Again,
as in California, the role of sea urchin grazing in regulating Macrocystis populations in southern
Chile is more controversial. Initial experimental results detected no effect of Loxechinus albus
grazing on Macrocystis populations in the Beagle Channel (Castilla & Moreno 1982). On the other
hand, Dayton (1985b) argued that Loxechinus albus grazing should significantly affect Macrocystis
abundance along the protected coast of southern Chile where large asteroid predators could serve
as a controlling factor and further suggested that the results of Castilla & Moreno (1982) were only
relevant to the southernmost subpolar area (Beagle Channel). Additionally, density of an annual
Macrocystis population in the archipelago region of southern Chile was significantly reduced from 24
to 2 sporophytes m−2 when Loxechinus albus densities exceeded 20 m−2 (Buschmann et al. 2004b).
These contradictions have developed into an unresolved controversy about the ecological role of sea
urchins in structuring Chilean Macrocystis populations (Vásquez & Buschmann 1997).
Based on these numerous field studies spanning the global range of Macrocystis, simple trophic
cascades do not seem to exist in Macrocystis-based systems. Various instances of overgrazing have
been described (Steneck et al. 2002) but they are generally short-lived, observed at local scales and
are often the result of overgrazing by particular trophic groups (e.g., sea urchins and amphipods).
Nevertheless, due to the high diversity and productivity of Macrocystis-based food webs (Rosenthal
et al. 1974, Pearse & Hines 1976, Castilla 1985, Foster & Schiel 1985, Graham 2004, Graham et al.
2007), these rare overgrazing events can have conspicuous ecological consequences (e.g., Graham
2004). The question still remains, however, as to why overgrazing is less frequent in Macrocystis-
based systems than in other kelp-based systems (e.g., Aleutians, North Atlantic, Japan; Steneck
et al. 2002).
Three striking features are common to Macrocystis systems worldwide and may be important
in buffering Macrocystis communities from overexploitation. The first is that all Macrocystis-based
food webs are relatively diverse. Such high diversity, especially when it occurs at higher-order
trophic levels, may provide a wider range of trophic interactions than less-diverse systems, mini-
mising the impact of grazing by any given herbivore species. Such ecological effects of high
diversity are supported by the field and experimental mesocosm studies of Byrnes et al. (2006),
who found that increased predator diversity decreased the impact of an assemblage of grazers on
Macrocystis biomass. In addition to the highly diverse systems of California, Beckley & Branch
(1992) enumerated 200+ taxa in a Macrocystis system at the Prince Edwards Islands. Castilla (1985)
identified 30+ herbivores and primary predators for the Macrocystis-based food web in the Beagle
Channel, culminating with the generalist asteroid Cosmarestias lurida; Adami & Gordillo (1999)
observed a similar system on the other side of the channel, although Loxechinus albus was
conspicuously absent. Vásquez et al. (1998) observed similar trophic diversity in the Macrocystis
system of northern Chile, which can include the Southern Hemisphere sea otter Lontra felina.
Although L. felina does not feed on sea urchins (Ebensperger & Botto-Mahan 1997, Villegas 2002),
these sea otters do forage on fishes, crustaceans and molluscs and may represent a diversifying
component in these systems (Castilla & Bahamondes 1979). A poleward decrease in the diversity
of Macrocystis systems appears to be present in both the Northern (Graham 2004, Graham et al.
2007) and Southern Hemispheres (Castilla 1985, Vásquez et al. 1998). The second commonality
69
among Macrocystis systems is that they are all imbedded within high-productivity systems necessary
to support Macrocystis survival, growth and reproduction. Therefore, the inherently high delivery
of nutrients to global Macrocystis populations may simply override consumptive processes in
regulating community structure and ecosystem processes over broad temporal and spatial scales.
Finally, sea urchin recruitment appears to be more variable in space and time at low latitudes
compared with high latitudes (Castilla & Moreno 1982, Foster & Schiel 1988, Buschmann et al.
2004b, Vega et al. 2005), potentially destabilising sea urchin population dynamics and decreasing
the likelihood of large-scale sea urchin population explosions (Foster & Schiel 1988).
Despite the numerous trophic studies of kelp forest organisms, however, there is a dearth of
research on communitywide patterns of energy flow. Stable isotope methods have demonstrated
the important role of detrital pathways in Macrocystis forests (e.g., Kaehler et al. 2000) and other
systems (Duggins et al. 1989, Bustamante & Branch 1996, Fredriksen 2003). Macrocystis produc-
tivity is also exported to other systems (e.g., sandy beaches, deep-sea basins, coastal islands) where
it may contribute greatly as an allochthonous energy source (Lavoie 1985, Inglis 1989, Vetter 1994,
Harrold et al. 1998, Orr et al. 2005). The trophic consequences of Macrocystis production, however,
have rarely been considered beyond the finite boundaries of the kelp forest.
Clearly, Macrocystis systems are energy rich. Trophic interactions among kelp forest organisms
can be conspicuous and are interesting avenues for ecological research. Yet, the emerging pattern
over the last 150 yr of research is that at the community or ecosystem level, the diversity and
productivity of Macrocystis systems are driven primarily by oceanographic processes that regulate
the distribution, abundance and standing stock of the main foundation species, Macrocystis. It has
recently been proposed, however, that the primary structural force for Macrocystis-based systems
in southern California (Channel Islands National Park) is ‘top-down’ consumption (Halpern et al.
2006). Halpern et al. (2006) used satellite-derived chlorophyll-a data to estimate nutrient delivery
to kelp beds as a proxy for ‘bottom-up’ processes. The effect of nutrients on algal abundance
(primarily that of Macrocystis) was then determined to be significantly lower than consumptive
effects. Off-shore chlorophyll-a concentrations, however, are not indicative of processes acting in
the near shore (Blanchette et al. 2006) and nutrient delivery to off-shore plankton assemblages and
near-shore kelp beds are two fundamentally different and negatively correlated processes (Broitman
& Kinlan 2006). Additionally, it is well established that variability in Macrocystis sporophyte
density (the abundance variable used by Halpern et al. 2006) is driven primarily by self-thinning
and is unrelated to nutrient supply (North 1994), whereas nutrient supply and Macrocystis biomass
are tightly coupled (North 1994, Tegner et al. 1996, 1997, Dayton et al. 1999). It would be inter-
esting to know whether Halpern et al. (2006) would have obtained different results if they had used
Macrocystis canopy biomass data available for the same region (Reed et al. 2006) and conducted
their study beyond 1999–2002, which was the most ‘nutrient benign’ period of the last 50 yr. In
fact, their study period did not include any of the conspicuous El Niño or La Niña events known
to drive maxima and minima in community structure and energy flow within these systems (Dayton
et al. 1999, Edwards 2004).
A final concern with the approach of Halpern et al. (2006) is the inability of their correlative
analyses to disentangle the confounding effects of habitat versus trophic associations. For example,
one of their four conspicuous species that was correlated with Macrocystis abundance, and thus
identified as a key consumer, was the striped surfperch (Embiotoca lateralis). Previous studies have
repeatedly observed a negative association between E. lateralis and Macrocystis (Ebeling & Laur
1985, Holbrook et al. 1990). The mechanism underlying the association, however, is not a top-
down trophic interaction but rather the negative effect of Macrocystis surface canopies on preferred
foraging habitat (foliose algae) of Embiotoca lateralis. Another of the conspicuous species, the
scavenger Kelletia kelletii, was found previously to be associated with sea urchin barrens rather
than kelp forests (Behrens & Lafferty 2004), the exact opposite pattern from that predicted by the
70
top-down hypothesis. The confounding nature of habitat versus trophic interactions in driving kelp
forest associations is probably ubiquitous among Macrocystis systems due to this species simulta-
neous provision of primary habitat and energy throughout much of its range (see ‘Macrocystis as
a foundation species,’ p. 62). This criticism of the results of Halpern et al. (2006) does not mean
that predation is unimportant in regulating the structure and dynamics of Macrocystis systems, but
simply argues for greater caution when using correlative data to understand regulatory processes
in this complex system.
Community consequences of climate change and kelp forest exploitation

Climate change and human exploitation can affect the diversity and productivity of Macrocystis
systems either by indirect modification of Macrocystis distribution, abundance and productivity or
by directly modifying distribution, abundance and productivity of the flora and fauna that inhabit
Macrocystis forests. Macrocystis productivity and distributional limits are largely constrained by
environmental processes (see ‘Organismal biology’ section, p. 42). Studies of environmental control
on Macrocystis systems, however, have been limited entirely to ecological timescales (e.g., Dayton
et al. 1999). Over periods of years to decades, temporal changes in nutrient availability (as measured
through sea-surface temperature proxies), sedimentation and substratum composition, storms and
light have all been shown to modify the living space and carrying capacity of Macrocystis (e.g.,
North & Schaeffer 1964, Zimmerman & Robertson 1985, Seymour et al. 1989, North 1994, Tegner
et al. 1996, 1997, Dayton et al. 1999, Edwards 2004). Such responses to short-term climate change
have typically been associated with ENSOs (4- to 7-yr frequency; Dayton et al. 1999, Edwards
2004) and cycles in the Pacific Decadal Oscillation (PDO, 10- to 20-yr frequency; Dayton et al.
1999). In California, strong ENSOs affect Macrocystis populations in two primary ways: nutrient
stress associated with deepening of the thermocline and destructive storm waves. The relative
impacts of each of these processes, however, vary latitudinally (Edwards 2004). Conspicuous
second-order community responses often result, for example, from reduction in the availability of
Macrocystis standing stock and drift and subsequent overgrazing by sea urchins or crustaceans
(Ebeling et al. 1985, Harrold & Reed 1985, Tegner & Dayton 1987, Graham 2002, Behrens &
Lafferty 2004). Ecologically important echinoderms (e.g., sea urchins and seastars) often suffer
mass mortalities that may also be associated with ENSOs (Tegner & Dayton 1987, Dayton &
Tegner 1989, Dayton et al. 1992, Behrens & Lafferty 2004, Lafferty 2004). These high-frequency
ENSO cycles are overlaid on longer-frequency PDO cycles, with warm PDO periods exacerbating
ENSO cycles (Dayton et al. 1999); the two most destructive ENSOs on record (1982–1983 and
1997–1998) occurred during the most recent warm PDO period. Nevertheless, the most well-
established effect of PDO cycling on Macrocystis systems is the correlation between larger Mac-
rocystis sporophyte sizes during PDO cold periods relative to warm periods (Tegner et al. 1996,
1997).
The importance of short-term oceanographic phenomena (e.g., ENSO) in regulating other
Macrocystis systems is essentially unknown. During the 1997–1998 El Niño, northern Chilean
Macrocystis populations increased, while black sea urchins (Tetrapygus niger) decreased (Vega
et al. 2005), a pattern opposite to that observed in southern California. The 1997–1998 El Niño
devastated Macrocystis populations in central Perú, decreasing sporophyte density and the diversity
of associated species (Lleellish et al. 2001). The subsequent 1998–1999 La Niña, however, triggered
high Tetrapygus niger recruitment and a significant increase in T. niger adult populations (Vásquez
et al. 2006), which corresponded with a crash in Macrocystis populations (Vega et al. 2005). The
1997–1998 El Niño also affected northern Chilean asteroid populations. Luidia and Meyenaster
are considered to be top predators in littoral benthic food chains of northern Chile, and both species
prey upon Heliaster and Stichaster (Viviani 1979). Luidia and Meyenaster coexist and restrict the
71
bathymetric distribution of Stichaster and Heliaster in the intertidal and subtidal zones (Viviani
1979). Meyenaster and Luidia decreased significantly within Macrocystis populations during the
1997–1998 El Niño, potentially migrating to deeper water, whereas Heliaster and Stichaster
increased in abundance during the same period (Vásquez et al. 2006). It remains to be determined
whether the ENSO-driven decreases in Luidia and Meyenaster abundances, both important predators
on Tetrapygus (Viviani 1979, Vásquez 1993, Vásquez & Buschmann 1997), were the ultimate
causes of the 1998–1999 T. niger population explosion (Vega et al. 2005, Vásquez et al. 2006).
Although nutrient deprivation is the most conspicuous intra- and interdecadal oceanographic
stressor on Macrocystis physiology and survival, it has recently been shown that temperature shifts
alone can have rapid impacts on Macrocystis systems from the organismal to community levels.
Schiel et al. (2004) used an 18-yr dataset to study changes in the structure of a local Nereocystis
kelp bed after 10 yr of increased ocean temperature (+3.5°C) due to the thermal outfall of a power-
generating station. Similar to the changes observed following deforestation in southern California
(Graham 2004), Schiel et al. (2004) detected significant communitywide changes in 150 species
of algae and invertebrates since the initiation of the thermal outfall. These community changes,
however, were not consistent with a northern shift in the distribution of southern species, but rather
a shift in the dominant canopy-forming kelp from Nereocystis to Macrocystis, and the potential
shading effect of the Macrocystis surface canopy. These data demonstrate the difficulty in disen-
tangling the direct effect of climate change on giant kelp communities from the indirect effect of
climate change on the distribution, abundance and productivity of key habitat-forming and energy-
producing species, like Macrocystis.
Studies of the effects of natural and anthropogenic climate change on Macrocystis systems
have been limited to the last few decades. The frequency and severity of ENSOs have been highly
variable over geological timescales (Rosenthal & Broccoli 2004) and it has been suggested that
their frequency is increasing (Diaz et al. 2001). Still, the ecological consequences of such long-
term climate change to Macrocystis systems seem obvious; Schimmelmann & Tegner (1991)
detected an ENSO signal in the flux of Macrocystis-derived organic carbon to the floor of the Santa
Barbara Basin over 1500 yr. Less obvious, however, are interactions between long-term changes
in ocean temperature, near-shore sedimentation, light and sea level that are driven by glacial-
interglacial cycling (Graham et al. 2003). Macrocystis has limited depth, substratum composition
and nutrient ranges, and ice age redistribution and modification of environmental conditions may
have had massive impacts on Macrocystis distribution, abundance and productivity. For example,
late-Quaternary sea-level rise probably led to large changes in inhabitable Macrocystis reef area
around the Californian Channel Islands and mainland as broad near-shore rocky platforms became
exposed, shrank and even fragmented (Graham et al. 2003, Kinlan et al. 2005). A recent study
predicted that southern Californian Macrocystis kelp forest area and biomass increased up to 3-fold
from the last glacial maximum to the mid-Holocene, but then rapidly declined by 40–70% during
the late Holocene to current area and biomass levels (M.H. Graham, B.P. Kinlan, R.K. Grossberg,
unpublished data). Furthermore, the early Holocene peak in Macrocystis distribution and abundance
coincided with highly productive palaeo-oceanographic conditions, probably yielding a subsequent
peak in Macrocystis productivity during that period. This shift overlapped with conspicuous changes
in total biomass of kelp-associated species, such as abalone, sea urchins and turban snails in native
American shell middens on the Channel Islands (Erlandson et al. 2005). The community and
ecosystem consequences of such long-term climate change on Macrocystis systems can be predicted
but critical tests of such predictions will require application of contemporary palaeo-ecological
tools (e.g., stable isotopes) because Macrocystis sporophytes do not fossilise (Graham et al. 2003).
Poor strategies of sewage discharge in the 1950s and 1960s were associated with the decimation
of a few very large Macrocystis forests in southern California (North & Schaeffer 1964, North &
Hubbs 1968, North 1971, Tegner & Dayton 1991). Stringent regulations, however, quickly remedied
72
the impacts. Tegner et al. (1995b) later found that nitrogenous wastes originating from breakage
in sewage outfalls can actually have positive effects on Macrocystis recruitment, especially during
periods of nutrient deprivation. It was also noted by Dawson et al. (1960) that an oil spill in Baja
California, Mexico, had no direct impacts on Macrocystis physiology, yet positively affected
Macrocystis survival by causing high local mortality of sea urchins. Effects of other pollutants,
such as some metals and aqueous petroleum waste, on Macrocystis microscopic stages can inhibit
microtubule dynamics, DNA replication, photosynthetic processes and overall physiology (Ander-
son et al. 1990, Garman et al. 1994, 1995, Reed & Lewis 1994). Despite these localised impacts,
there is little evidence that chemical pollution currently restricts Macrocystis distribution, abundance
and productivity over broad spatial and temporal scales.
Finally, Macrocystis systems have been subjected to long-term anthropogenic exploitation,
spanning a period of at least 11,000 yr (Erlandson et al. 2005). Recent attention has focused on
direct exploitation of Macrocystis populations and Macrocystis-associated organisms, especially in
southern California where Macrocystis has been harvested for algin extraction since the 1920s
(North 1994). Californian harvests are limited to the upper 1–2 m of the water column and have
been shown to have minimal impacts on sporophyte survival (see p. 56). Indeed, while there is
considerable temporal variability in Macrocystis populations due to physical and biological factors,
the long-term stability of the Macrocystis harvest suggests that it is one of the best-managed marine
harvests of wild populations worldwide (Dayton et al. 1998). Nevertheless, in southern Chile,
Macrocystis is harvested by abalone farmers who require biomass all year round and Macrocystis
cultivation is now required to offset heavy exploitation of natural Macrocystis populations (Gutierrez
et al. 2006). Due to the patchy distribution of integrifolia-form populations in northern Chile,
Macrocystis harvesting near abalone farms has had a great impact on the dynamics of local
Macrocystis populations, with subsequent effects on Macrocystis-associated communities (J.A.
Vásquez, unpublished data).
Although Macrocystis populations themselves appear to be relatively immune to episodic
harvesting of the surface canopy, Macrocystis-associated organisms are not. Overfishing has resulted
in virtual elimination of large predators in southern California Macrocystis forests (Dayton et al.
1998). The ecological impacts of overfishing on Macrocystis populations are unclear because some
correlative studies suggest cascading impacts whereas others do not (Foster & Schiel 1988, Dayton
et al. 1998, Steneck et al. 2002, Behrens & Lafferty 2004, Lafferty 2004, Graham et al. 2007).
Although predators may or may not be more common within marine reserves (Paddack & Estes
2000, Behrens & Lafferty 2004, Lafferty 2004), predators within reserves are typically larger in
size (Paddack & Estes 2000). Again, the problem lies in deciphering the various types and strengths
of species interactions operating in Macrocystis forests (e.g., Macrocystis-derived habitat and energy
provision compared with predation). One thing is clear, however, despite the ubiquitous role of
Macrocystis-derived habitat and energy provision in enhancing kelp forest diversity and productivity
worldwide, kelp forest organisms cannot survive targeted exploitation over large temporal and
spatial scales (Dayton et al. 1998).
Conclusion
The global scientific literature indicates that Macrocystis is an important provider of habitat and
energy to its associated communities wherever it is present. It is also clear that, despite non-trivial
gene flow among global Macrocystis populations, Macrocystis morphology and physiology are
highly variable in response to the environmental conditions within which sporophytes recruit, grow
and reproduce. These conspicuous ecotypic differences have generally led researchers to study
Macrocystis population dynamics and community interactions from a regional perspective. Patterns
observed for some large conspicuous giant kelp forests in southern California have subsequently
73
dominated the literature and become the paradigms against which the ecologies of other Macrocystis
systems are compared. When viewed from a global perspective, however, regional differences in
the results of prior descriptive and experimental studies can be reconciled by an appreciation of
great plasticity in Macrocystis form and function. The origin, nature and potential restriction of such
plasticity to Macrocystis are appealing paths for future research.
Acknowledgements
We greatly appreciate the support of our funding agencies: J.A.V. and A.H.B. acknowledge
FONDECYT (grants 1010706, 1000044 and 1040425) and the Universidad de Los Lagos; M.H.G.
acknowledges the National Science Foundation (NSF 0351778 and 0407937), San Jose State
University, and the Moss Landing Marine Laboratories. We also acknowledge Alonso Vega, Mariam
Hernández, Pirjo Huovenin, René Espinoza, Lara Ferry-Graham, David Schiel, Sean Connell, Louis
Druehl and Michael Foster for various levels of support and comments during production of this
review.
References
Abbott, I.A. & Hollenberg, G.J. 1976. Marine Algae of California. Stanford, California: Stanford University
Press.
Adami, M.L. & Gordillo, S. 1999. Structure and dynamics of the biota associated with Macrocystis pyrifera
(Phaeophyta) from the Beagle Channel, Tierra del Fuego. Scientia Marina 63, 183–191.
Aguilar-Rosas, L.E., Aguilar-Rosas, R., Marcos-Ramirez, R., Caceres-Rubio, C.F. & McPeak, R.H. 2003.
New record of Macrocystis laevis (Laminariales, Phaeophyta) on the Pacific coast of Chile. Proceed-
ings of the International Seaweed Symposium 17, 337–340.
Amsler, C.D. & Neushul, M. 1989. Chemotactic effects of nutrients on spores of the kelps Macrocystis pyrifera
and Pterygophora californica. Marine Biology 102, 557–564.
Amsler, C.D. & Neushul, M. 1990. Nutrient stimulation of spore settlement in the kelps Pterygophora
californica and Macrocystis pyrifera. Marine Biology 107, 297–304.
Amsler, C.D., Reed, D.C. & Neushul, M. 1992. The microclimate inhabited by macroalgal propagules. British
Phycological Journal 27, 253–270.
Anderson, B.S., Hunt, J.W., Turpen, S.L., Coulon, A.R. & Martin, M. 1990. Copper toxicity to microscopic
stages of giant kelp Macrocystis pyrifera: interpopulation comparisons and temporal variability.
Marine Ecology Progress Series 68, 147–156.
Anderson, E.K. & North, W.J. 1966. In situ studies of spore production and dispersal in the giant kelp
Macrocystis pyrifera. Proceedings of the International Seaweed Symposium 5, 73–86.
Anderson, T.W. 1994. Role of macroalgal structure in the distribution and abundance of a temperate reef fish.
Anderson, T.W. 2001. Predator responses, prey refuges, and density-dependent mortality of a marine fish.
Ecology 82, 245–257.
Andrews, N.L. 1945. The kelp beds of the Monterey region. Ecology 26, 24–37.
Arnold, K.E. & Manley, S.L. 1985. Carbon allocation in Macrocystis pyrifera (Phaeophyta): intrinsic variability
in photosynthesis and respiration. Journal of Phycology 21, 154–167.
Asensi, A., Beucher, M. & Delepine, R. 1981. Études sur la croissance et le développement de Macrocystis
pyrifera (L.) C. Ag. aux Iles Kerguelen. Proceedings of the International Seaweed Symposium 8, 278–289.
Attwood, C.G., Lucas, M.I., Probyn, T.A., McQuaid, C.D. & Fielding, P.J. 1991. Production and standing
stocks of the kelp Macrocystis laevis Hay at the Prince Edward Islands, Subantarctic. Polar Biology
11, 129–133.
Augenstein, E.W., Stow, D.A. & Hope, A.S. 1991. Evaluation of SPOT HRV-XS data for kelp resource
inventories. Photogrammetric Engineering and Remote Sensing 57, 501–509.
74
Avila, M., Hoffmann, A.J. & Santelices, B. 1985. Interactions of temperature, photon flux density and photo-
period on the development of microscopic stages of Lessonia nigrescens (Phaeophyta, Laminariales).
Revista Chilena de Historia Natural 58, 71–82.
Baardseth, E. 1941. The marine algae of Tristan da Cunha. Results of the Norwegian Scientific Expedition to
Tristan da Cunha 1937–1938 9, 1–160.
Barilotti, D.C., McPeak, R.H. & Dayton, P.K. 1985. Experimental studies on the effects of commercial kelp
harvesting in central and southern California Macrocystis pyrifera kelp beds. California Fish and
Game 71, 4–20.
Barrales, H.L. & Lobban, C.S. 1975. The comparative ecology of Macrocystis pyrifera, with emphasis on the
forests of Chubut, Argentina. Journal of Ecology 63, 657–677.
Beckley, L.E. & Branch, G.M. 1992. A quantitative scuba-diving survey of the sublittoral macrobenthos at
subantarctic Marion Island. Polar Biology 11, 553–563.
Behrens, M.D. & Lafferty, K.D. 2004. Effects of marine reserves and urchin disease on southern Californian
rocky reef communities. Marine Ecology Progress Series 279, 129–139.
Belsher, T. & Mouchot, M.C. 1992. Use of satellite imagery in management of giant kelp resources Morbihan
Gulf, Kerguelen Archipelago. Oceanologica Acta 15, 297–307.
Bernstein, B.L. & Jung, N. 1979. Selective pressures and coevolution in a kelp canopy community in southern
California. Ecological Monographs 49, 335–355.
Blanchette, C.A., Broitman, B.R. & Gaines S.D. 2006. Intertidal community structure and oceanographic
patterns around Santa Cruz Island, California, U.S.A. Marine Biology 149, 689–701.
Bray, R.N. & Ebeling, A.W. 1975. Food, activity, and habitat of three ‘picker-type’ microcarnivorous fishes
in the kelp forests off Santa Barbara, California. Fishery Bulletin 73, 815–829.
Broitman B.R. & Kinlan B.P. 2006. Spatial scales of benthic and pelagic producer biomass in a coastal
upwelling ecosystem. Marine Ecology Progress Series 327, 15–25.
Brostoff, W.N. 1988. Taxonomic studies of Macrocystis pyrifera (L) C. Agardh (Phaeophyta) in southern
California: holdfasts and basal stipes. Aquatic Botany 31, 289–305.
Brown, M.T., Nyman, M.A., Keogh, J.A. & Chin, N.K.M. 1997. Seasonal growth of the giant kelp Macrocystis
pyrifera in New Zealand. Marine Biology 129, 417–424.
Burgman, M.A. & Gerard, V.A. 1990. A stage-structured, stochastic population model for the giant kelp
Macrocystis pyrifera. Marine Biology 105, 15–23.
Buschmann, A.H. 1992. Algal communities of a wave-protected intertidal rocky shore in southern Chile. In
Coastal Plant Communities of Latin America, U. Seeliger (ed.). Miami, Florida: Academic Press, 91–104.
Buschmann, A.H., García, C., Espinoza, R., Filún, L. & Vásquez, J.A. 2004b. Sea urchin (Loxechinus albus)
and kelp (Macrocystis pyrifera) interaction in protected areas in southern Chile. In Sea Urchin Biology,
J. Lawrence & O. Guzmán (eds). Lancaster, Pennsylvania: DEStech Publications, 120–130.
Buschmann, A.H., Moreno, C.A., Vásquez, J.A. & Hernández-González, M.C. 2006. Population and repro-
duction strategies of Macrocystis pyrifera (Phaeophyta) in southern Chile. Journal of Applied Phy-
cology 18, 575–582.
Buschmann, A.H., Vásquez, J.A., Osorio, P., Reyes, E., Filún, L., Hernández-González, M.C. & Vega, A.
2004a. The effect of water movement, temperature and salinity on abundance and reproductive patterns
of Macrocystis spp. (Phaeophyta) at different latitudes in Chile. Marine Biology 145, 849–862.
Bustamante, R.H. & Branch, G.M. 1996. The dependence of intertidal consumers on kelp-derived organic
matter on the west coast of South Africa. Journal of Experimental Marine Biology and Ecology 196,
1–28.
Byrnes, J., Stachowicz, J.J., Hultgren, K.M., Hughes, A.R., Olyarnik, S.V. & Thornber, C.S. 2006. Predator
diversity strengthens trophic cascades in kelp forests by modifying herbivore behaviour. Ecology
Letters 9, 61–71.
Camus, P.A. 1994. Recruitment of the intertidal kelp Lessonia nigrescens Bory in northern Chile: successional
constraints and opportunities. Journal of Experimental Marine Biology and Ecology 184, 171–181.
Carr, M.H. 1989. Effects of macroalgal assemblages on the recruitment of temperate zone reef fishes. Journal
of Experimental Marine Biology and Ecology 126, 59–76.
Carr, M.H. 1991. Habitat selection and recruitment of an assemblage of temperate zone reef fishes. Journal
75
Carr, M.H. 1994. Effects of macroalgal dynamics on recruitment of a temperate reef fish. Ecology 75,
1320–1333.
Castilla, J.C. 1985. Food webs and functional aspects of the kelp, Macrocystis pyrifera, community in the
Beagle Channel, Chile. In Antarctic Nutrient Cycles and Food Webs, W.R. Siegfried et al. (eds).
Heidelberg, Germany: Springer-Verlag, 407–414.
Castilla, J.C. & Bahamondes, I. 1979. Observaciones conductuales y ecologicas sobre Lutra felina en las
zonas central y centro-norte de Chile. Archivos de Biologia y Medicina Experimentales 12, 119–132.
Castilla, J.C. & Moreno, C.A. 1982. Sea urchins and Macrocystis pyrifera: Experimental test of their ecological
relations in southern Chile. In Proceedings of the International Echinoderm Conference, J.M.
Lawrence (ed.). Rotterdam: A.A. Balkema, 257–263.
Chamberlain, Y.M. 1965. Marine algae of Gough Island. Bulletin of the British Museum (Natural History)
Botany 3, 173–232.
Chapman, A.R.O. & Craigie, J.S. 1977. Seasonal growth in Laminaria longicruris: relations with dissolved
inorganic nutrients and internal reserves of nitrogen. Marine Biology 40, 197–205.
Chin, N.K.M., Brown, M.T. & Heads, M.J. 1991. The biogeography of Lessoniaceae, with special reference
to Macrocystis C Agardh (Phaeophyta, Laminariales). Hydrobiologia 215, 1–11.
Clark, R.P., Edwards, M.S. & Foster, M.S. 2004. Effects of shade from multiple kelp canopies on an understory
algal assemblage. Marine Ecology Progress Series 267, 107–119.
Clarke, W.D. 1971. Mysids of the southern kelp region. Nova Hedwigia 32, 369–380.
Cowen, R.K. 1983. The effect of sheephead (Semicossyphus pulcher) predation on Red Sea urchin (Strongy-
locentrotus franciscanus) populations: an experimental analysis. Oecologia 58, 249–255.
Cowen, R.K. 1986. Site-specific differences in the feeding ecology of the California sheephead, Semicossyphus
pulcher (Labridae). Environmental Biology of Fishes 16, 193–203.
Cowen, R.K., Agegian, C.R. & Foster, M.S. 1982. The maintenance of community structure in a central
California giant kelp forest. Journal of Experimental Marine Biology and Ecology 64, 189–201.
Coyer, J.A. 1985. The invertebrate assemblage associated with the giant kelp, Macrocystis pyrifera, at Santa
Catalina Island, California: a general description with emphasis on amphipods, copepods, mysids and
shrimps. Fishery Bulletin 82, 55–66.
Coyer, J.A. 1987. The mollusk assemblage associated with fronds of giant kelp (Macrocystis pyrifera) off
Santa Catalina Island, California. Bulletin of the Southern California Academy of Sciences 85, 129–138.
Coyer, J.A., Smith, G.J. & Andersen, R.A. 2001. Evolution of Macrocystis spp. (Phaeophyceae) as determined
by ITS1 and ITS2 sequences. Journal of Phycology 37, 574–585.
Crandall, W.C. (ed.) 1915. Potash from kelp. U.S. Department of Agriculture Report 100, Part II, Washington,
D.C.
Cribb, A.B. 1954. Macrocystis pyrifera (L.) Ag. in Tasmanian waters. Australian Journal of Marine and
Freshwater Research 5, 1–34.
Darwin, C.R. 1839. Journal of Researches into the Geology and Natural History of the Various Countries
Visited by the H.M.S. Beagle. London, England: Henry Colburn.
Dawson, E.Y., Neushul, M. & Wildman, R.D. 1960. Seaweeds associated with kelp beds along southern
California and northwestern México. Pacific Naturalist 1, 2–81.
Dayton, P.K. 1985a. Ecology of kelp communities. Annual Review of Ecology and Systematics 16, 215–245.
Dayton, P.K. 1985b. The structure and regulation of some South American kelp communities. Ecological
Monographs 55, 447–468.
Dayton, P.K., Currie, V., Gerrodette, T., Keller, B.D., Rosenthal, R.J. & Van Tresca, D. 1984. Patch dynamics
and stability of some California kelp communities. Ecological Monographs 54, 253–289.
Dayton, P.K., Rosenthal, R.J. & Mahan, L.C. 1973. Kelp communities in the Chilean Archipelago: R/V Hero
cruise 1972–1975. Antarctic Journal of the United States 8, 34–35.
Dayton, P.K. & Tegner, M.J. 1984. Catastrophic storms, El Niño, and patch stability in a southern California
kelp community. Science 224, 283–285.
Dayton, P.K. & Tegner, M.J. 1989. Bottoms beneath troubled water, benthic impacts of the 1982–1984 El
Niño in the temperate zone. In Global Ecological Consequences of the 1982–1983 El Niño-Southern
Oscillation, P.W. Glynn (ed.). Elsevier Oceanography Series 52, Amsterdam, Netherlands: Elsevier
Science.
76
Dayton, P.K., Tegner, M.J., Edwards, P.B. & Riser, K.L. 1998. Sliding baselines, ghosts, and reduced expec-
tations in kelp forest communities. Ecological Applications 8, 309–322.
Dayton, P.K., Tegner, M.J., Edwards, P.B. & Riser, K.L. 1999. Temporal and spatial scales of kelp demography:
the role of oceanographic climate. Ecological Monographs 69, 219–250.
Dayton, P.K., Tegner, M.J., Parnell, P.E. & Edwards, P.B. 1992. Temporal and spatial patterns of disturbance
and recovery in a kelp forest community. Ecological Monographs 62, 421–445.
Dean, T.A. & Jacobsen, F.R. 1984. Growth of juvenile Macrocystis pyrifera (Laminariales) in relation to
environmental factors. Marine Biology 83, 301–311.
Dean, T.A. & Jacobsen, F.R. 1986. Nutrient-limited growth of juvenile kelp Macrocystis during the 1982–1984
‘El Niño’ in southern California. Marine Biology 90, 597–601.
Dean, T.A., Jacobsen, F.R., Thies, K. & Lagos, S.L., 1988. Differential effects of grazing by white sea urchins
on recruitment of brown algae. Marine Ecology Progress Series 48, 99–102.
Dean, T.A., Schroeter, S.C. & Dixon, J.D. 1984. Effects of grazing by two species of sea urchins (Strongy-
locentrotus franciscanus and Lytechinus anamesus) on recruitment and survival of two species of kelp
(Macrocystis pyrifera and Pterygophora californica). Marine Biology 78, 301–313.
Dean, T.A., Thies, K. & Lagos, S.L. 1989. Survival of juvenile giant kelp: the effects of demographic factors,
competitors, and grazers. Ecology 70, 483–495.
Delépine, R. 1966. La végétation marine dans l’Antarctique de l’ouest comparée à celle des Îles Australes
Francaises: conséquences biogéographiques. Comptes Rendus des Seances de la Société de Biogéog-
raphie 374, 52–68.
Delille, B., Delille, D., Fiala, M., Prevost, C. & Frankignoulle, M. 2000. Seasonal changes of pCO2 over a
subantarctic Macrocystis kelp bed. Polar Biology 23, 706–716.
DeMartini, E.E. & Roberts, D.A. 1990. Effects of giant kelp Macrocystis on the density and abundance of
fishes in a cobble-bottom kelp forest. Bulletin of Marine Science 46, 287–300.
Devinny, J.S. & Volse, L.A. 1978. Effects of sediments on the development of Macrocystis pyrifera gameto-
phytes. Marine Biology 48, 343–348.
DeWreede, R.E. & Klinger, T. 1988. Reproductive strategies in algae. In Plant Reproductive Ecology, J.L.
Doust & L.L. Doust (eds). New York: Oxford University Press, 267–284.
Deysher, L.E. 1993. Evaluation of remote sensing techniques for monitoring giant kelp populations. Hydro-
biologia 260/261, 307–312.
Deysher, L.E. & Dean, T.A. 1984. Critical irradiance levels and the interactive effects of quantum irradiance
and dose on gametogenesis in the giant kelp Macrocystis pyrifera. Journal of Phycology 20, 520–524.
Deysher, L.E. & Dean, T.A. 1986a. Interactive effects of light and temperature on sporophyte production in
the giant kelp Macrocystis pyrifera. Marine Biology 93, 17–20.
Deysher, L.E. & Dean, T.A. 1986b. In situ recruitment of sporophytes of the giant kelp Macrocystis pyrifera
(L.) C.A. Agardh: effects of physical factors. Journal of Experimental Marine Biology and Ecology
103, 41–63.
Diaz, H.F., Hoerling, M.P. & Eischeid, J.K. 2001. ENSO variability, teleconnections and climate change.
International Journal of Climatology 21, 1845–1862.
Dixon, J., Schroeter, S.C. & Kastendiek, J. 1981. Effects of the encrusting bryozoan, Membranipora mem-
branacea, on the loss of blades and fronds by the giant kelp, Macrocystis pyrifera (Laminariales).
Journal of Phycology 17, 341–345.
Donnellan, M.D. 2004. Spatial and temporal variability of kelp forest canopies in central California. M.S.
thesis, San Jose State University, San Jose, California.
Downes, B.J. & Keough, M.J. 1998. Scaling of colonization processes in streams: parallels and lessons from
marine hard substrata. Australian Journal of Ecology 23, 8–26.
Druehl, L.D. 1978. The distribution of Macrocystis integrifolia in British Columbia as related to environmental
factors. Canadian Journal of Botany 56, 69–79.
Druehl, L.D. & Breen, P.A. 1986. Some ecological effects of harvesting Macrocystis integrifolia. Botanica
Marina 29, 97–103.
Druehl, L.D., Collins, J.D., Lane, C.D. & Saunders, G.W. 2005. An evaluation of methods used to assess
intergeneric hybridization in kelp using Pacific Laminariales (Phaeophyceae). Journal of Phycology
41, 250–262.
77
Druehl, L.D. & Kemp, L. 1982. Morphological and growth responses of geographically isolated Macrocystis
integrifolia populations when grown in a common environment. Canadian Journal of Botany 60,
1409–1413.
Druehl, L.D., Mayes, C., Tan, I.H. & Saunders, G.W. 1997. Molecular and morphological phylogenies of kelp
and associated brown algae. In Plant Systematics and Evolution Supplement 11: Origins of Algae and
their Plastids, D. Bhattacharya (ed.). Vienna: Springer-Verlag, 221–235.
Druehl, L.D., Robertson, B.R. & Button, D.K. 1989. Characterizing and sexing Laminarialean meiospores by
flow cytometry. Marine Biology 101, 451–456.
Druehl, L.D. & Wheeler, W.N. 1986. Population biology of Macrocystis integrifolia from British Columbia,
Canada. Marine Biology 90, 173–179.
Duggins, D.O., Simenstad, C.A. & Estes, J.A. 1989. Magnification of secondary production by kelp detritus
in coastal marine ecosystems. Science 245, 170–173.
Dunton, K.H. 1990. Growth and production in Laminaria solidungula: relation to continuous underwater light
levels in the Alaskan high Arctic. Marine Biology 106, 297–304.
Dunton, K.H. & Jodwalis, C.M. 1988. Photosynthetic performance of Laminaria solidungula measured in
situ in the Alaskan high Arctic. Marine Biology 98, 277–286.
Eardley, D.D., Sutton, C.W., Hempel, W.M., Reed, D.C. & Ebeling, A.W. 1990. Monoclonal antibodies specific
for sulfated polysaccharides on the surface of Macrocystis pyrifera (Phaeophyceae). Journal of
Phycology 26, 54–62.
Ebeling, A.W. & Laur, D.R. 1985. The influence of plant cover on surfperch abundance at an offshore temperate
reef. Environmental Biology of Fishes 12, 169–180.
Ebeling, A.W., Laur, D.R. & Rowley, R.J. 1985. Severe storm disturbances and reversal of community structure
in a southern California kelp forest. Marine Biology 84, 287–294.
Ebensperger, L.A. & Botto-Mahan, C. 1997. Use of habitat, size of prey, and food-niche relationships of two
sympatric otters in southernmost Chile. Journal of Mammalogy 78, 222–227.
Edgar, G.J. 1987. Dispersal of faunal and floral propagules associated with drifting Macrocystis pyrifera plants.
Marine Biology 95, 599–610.
Edwards, M.S. 1998. Effects of long-term kelp canopy exclusion on the abundance of the annual alga
Desmarestia ligulata (Light F). Journal of Experimental Marine Biology and Ecology 228, 309–326.
Edwards, M.S. 2004. Estimating scale-dependency in disturbance impacts: El Niños and giant kelp forests in
the northeast Pacific. Oecologia 138, 436–447.
Edwards, M.S. & Hernández-Carmona, G. 2005. Delayed recovery of giant kelp near its southern range limit
in the North Pacific following El Niño. Marine Biology 147, 273–279.
Erlandson, J.M., Rick, T.C., Estes, J.A., Graham, M.H., Braje, T.J. & Vellanoweth, R.L. 2005. Sea otters,
shellfish, and humans: a 10,000 year record from San Miguel island, California Proceedings of the
California Islands Symposium 6, 56–68.
Estes, J.A. & Duggins, D.O. 1995. Sea otters and kelp forests in Alaska: generality and variation in a community
ecological paradigm. Journal of Phycology 65, 75–100.
Estes, J.A. & Steinberg, P.D. 1988. Predation, herbivory, and kelp evolution. Paleobiology 14, 19–36.
Fain, S.R. & Murray, S.N. 1982. Effects of light and temperature on net photosynthesis and dark respiration
of gametophytes and embryonic sporophytes of Macrocystis pyrifera. Journal of Phycology 18, 92–98.
Feder, H.M., Turner, C.H. & Limbaugh, C. 1974. Observations on fishes associated with kelp beds in southern
California. California Department of Fish and Game, Fish Bulletin 160, 1–144.
Fosberg, F.R. 1929. Preliminary notes on the fauna of the giant kelp. Journal of Entolomological Zoology 21,
133–135.
Foster, M.S. 1982. The regulation of macroalgal associations in kelp forests. In Synthetic and Degradative
Processes in Marine Macrophytes, L. Srivastava (ed.). Berlin: Walter de Gruyter & Co., 185–205.
Foster, M.S. & Schiel, D.R. 1985. The ecology of giant kelp forests in California: a community profile. United
States Fish and Wildlife Service Biological Report 85, 1–152.
Foster, M.S. & Schiel, D.R. 1988. Kelp communities and sea otters: keystone species or just another brick in
the wall? In The Community Ecology of Sea Otters, G.R. VanBlaricom & J.A. Estes (eds). Berlin:
Springer-Verlag, 92–115.
78
Foster, M.S. & Schiel, D.R. 1992. Zonation, El Niño disturbance, and the dynamics of subtidal vegetation
along a 30 m depth gradient in two giant kelp forests. Proceedings of the International Temperate
Reef Symposium 2, 151–162.
Fredriksen, S. 2003. Food web studies in a Norwegian kelp forest based on stable isotope (d13C and d15N)
analysis. Marine Ecology Progress Series 260, 71–81.
Fritsch, F.E. 1945. The Structure and Reproduction of the Algae. Volume 2. Cambridge: Cambridge University
Press.
Gabrielson, P.W., Widdowson, T.B., Lindstrom, S.C., Hawkes, M.W. & Scagel, R.F. 2000. Keys to the Benthic
Marine Algae and Seagrasses of British Columbia, Southeast Alaska, Washington and Oregon: Phy-
cological Contribution #5. Vancouver: University of British Columbia Department of Botany.
Garman, G.D., Pillai, M.C. & Cherr, G.N. 1994. Inhibition of cellular events during algal gametophyte
development: effects of select metals and an aqueous petroleum waste. Aquatic Toxicology 28, 127–144.
Garman, G.D., Pillai, M.C., Goff, L.J. & Cherr, G.N. 1995. Nuclear events during early development in
Macrocystis pyrifera gametophytes and the temporal effects of a marine contaminant. Marine Biology
121, 355–362.
Gaylord, B., Reed, D.C., Raimondi, P.T., Washburn, L. & McLean, S.R. 2002. A physically based model of
macroalgal spore dispersal in the wave and current-dominated nearshore. Ecology 83, 1239–1251.
Gerard, V.A. 1976. Some aspects of material dynamics and energy flow in a kelp forest in Monterey Bay,
California. Ph.D. dissertation, University of California Santa Cruz, Santa Cruz, California.
Gerard, V.A. 1982. Growth and utilization of internal nitrogen reserves by the giant kelp Macrocystis pyrifera
in a low-nitrogen environment. Marine Biology 66, 27–35.
Gerard, V.A. & Kirkmann, H. 1984. Ecological observations on a branched, loose-lying form of Macrocystis
pyrifera (L) C. Agardh in New Zealand. Botanica Marina 27, 105–109.
Gerard, V.A. and Mann, K.H. 1979. Growth and production of Laminaria longicruris (Phaeophyta) populations
exposed to different intensities of water movement. Journal of Phycology 14, 195–198.
Ghelardi, R.J. 1971. The biology of giant kelp beds (Macrocystis) in California: species structure of the
holdfast community. Nova Hedwigia 32, 381–420.
Gonzalez-Fragoso, J., Ibarra-Obando, S.E. & North, W.J. 1991. Frond elongation rates of shallow water Macro-
cystis pyrifera (L.) Ag. in northern Baja California, México. Journal of Applied Phycology 3, 311–318.
Graham, M.H. 1996. Effect of high irradiance on recruitment of giant kelp Macrocystis (Phaeophyta) in
shallow water. Journal of Phycology 32, 903–906.
Graham, M.H. 1997. Factors determining the upper limit of giant kelp, Macrocystis pyrifera Agardh, along
the Monterey Peninsula, central California, U.S.A. Journal of Experimental Marine Biology and
Ecology 218, 127–149.
Graham, M.H. 1999. Identification of kelp zoospores from in situ plankton samples. Marine Biology 135,
709–720.
Graham, M.H. 2000. Planktonic patterns and processes in the giant kelp Macrocystis pyrifera. Ph.D. disser-
tation, University of California San Diego, La Jolla, California.
Graham, M.H. 2002. Prolonged reproductive consequences of short-term biomass loss in seaweeds. Marine
Biology 140, 901–911.
Graham, M.H. 2003. Coupling propagule output to supply at the edge and interior of a giant kelp forest.
Ecology 84, 1250–1264.
Graham, M.H. 2004. Effects of local deforestation on the diversity and structure of southern California giant
kelp forest food webs. Ecosystems 7, 341–357.
Graham, M.H., Dayton, P.K. & Erlandson, J.M. 2003. Ice ages and ecological transitions on temperate coasts.
Trends in Ecology and Evolution 18, 33–40.
Graham, M.H., Halpern, B.S. & Carr, M.H. 2007. Diversity and dynamics of Californian subtidal kelp forests.
In Food Webs and the Dynamics of Marine Benthic Ecosystems, T.R. McClanahan & G.R. Branch
(eds). Oxford: Oxford University Press, in press.
Graham, M.H., Harrold, C., Lisin, S., Light, K., Watanabe, J.M. & Foster, M.S. 1997. Population dynamics
of giant kelp Macrocystis pyrifera along a wave exposure gradient. Marine Ecology Progress Series
148, 269–279.
79
Graham, M.H. & Mitchell, B.G. 1999. Obtaining absorption spectra from individual macroalgal spores using
microphotometry. Hydrobiologia 398/399, 231–239.
Grua, P. 1964. Sur la structure des peuplements de Macrocystis pyrifera (L.) C. Ag. observés en plongée à
Kerguelen et Crozet. Comptes Rendus Hebdomadaires des Séances de l’Académie des Sciences Paris
259, 1541–1543.
Guiler, E.R. 1952. The intertidal ecology of Eaglehawk Neck area. Papers and Proceedings of the Royal
Society of Tasmania 86, 13–29.
Guiler, E.R. 1960. Notes on the intertidal ecology of Trial Harbour, Tasmania. Papers and Proceedings of the
Royal Society of Tasmania 94, 57–62.
Gutierrez, A., Correa, T., Muñoz, V., Santibañez, A., Marcos, R., Caceres, C. & Buschmann, A.H. 2006.
Farming of the giant kelp Macrocystis pyrifera in southern Chile for development of novel food
products. Journal of Applied Phycology 18, 259–267.
Hallacher, L.E. & Roberts, D.A. 1985. Differential utilization of space and food by the inshore rockfishes
(Scorpaenidae: Sebastes) of Carmel Bay, California [U.S.A.]. Environmental Biology of Fishes 12,
91–110.
Halpern, B.S., Cottenie, K. & Broitman, B.R. 2006. Strong top-down control in southern California kelp forest
ecosystems. Science 312, 1230–1232.
Harris, L.G., Ebeling, A.W., Laur, D.R. & Rowley, R.J. 1984. Community recovery after storm damage: a
case of facilitation in primary succession. Science 224, 1339–1338.
Harrold, C., Light, K. & Lisin, S. 1998. Organic enrichment of submarine-canyon and continental shelf benthic
communities by macroalgal drift imported from nearshore kelp forests. Limnology and Oceanography
43, 669–678.
Harrold, C. & Lisin, S. 1989. Radio-tracking rafts of giant kelp: Local production and regional transport.
Journal of Experimental Marine Biology and Ecology 130, 237–251.
Harrold, C. & Pearse, J.S. 1987. The ecological role of echinoderms in kelp forests. Echinoderm Studies 2,
137–233.
Harrold, C. & Reed, D.C. 1985. Food availability, sea urchin grazing, and kelp forest community structure.
Ecology 66, 1160–1169.
Hay, C.H. 1986. A new species of Macrocystis C. Ag. (Phaeophyta) from Marion Island, southern Indian
Ocean. Phycologia 25, 241–252.
Hay, C.H. 1990. The distribution of Macrocystis C. Ag. (Phaeophyta, Laminariales) as a biological indicator
of cool sea surface temperatures, with special reference to New Zealand water. Journal of the Royal
Society of New Zealand 20, 313–336.
Helmuth, B.S., Veit, R.R. & Holberton, R. 1994. Long-distance dispersal of subantarctic brooding bivalve
(Gaimardia trapesina) by kelp rafting. Marine Biology 120, 421–6.
Hempel, W.M., Sutton, C.W., Kaska, D., Ord, D.C., Reed, D.C., Laur, D.R., Ebeling, A.W. & Eardley, D.D.
1989. Purification of species-specific antibodies to carbohydrate components of Macrocystis pyrifera
(Phaeophyta). Journal of Phycology 25, 144–149.
Henry, E.C. & Cole, D.W. 1982. Ultrastructure of swarmers in the Laminariales (Phaeophyceae). I. Zoospores.
Hernández-Carmona, G. 1996. Frond elongation rates of Macrocystis pyrifera (L.) Ag. at Bahia Tortugas,
Baja California sur, México. Ciencias Marinas 22, 57–72.
Hernández-Carmona, G., Garcia, O., Robledo, D. & Foster, M.S. 2000. Restoration techniques for Macrocystis
pyrifera (Phaeophyceae) populations at the southern limit of their distribution in México. Botanica
Marina 43, 273–284.
Hernández-Carmona, G., Hughes, B. & Graham, M.H. 2006. Reproductive longevity of drifting kelp Macro-
cystis pyrifera (Phaeophyceae) in Monterey Bay, U.S.A. Journal of Phycology 42, 1199–1207.
Hernández-Carmona, G., Robledo, D. & Serviere-Zaragoza, E. 2001. Effect of nutrient availability on Macro-
cystis pyrifera recruitment and survival near its southern limit off Baja California. Botanica Marina
44, 221–229.
Hernández-Carmona, G., Rodriguez-Montesinos, Y.E., Casas-Valdez, M.M., Vilchis, M.A. & Sanchez-Rodriguez,
I. 1991. Evaluation of the beds of Macrocystis pyrifera (Phaeophyta, Laminariales) in the Baja California
peninsula, México. III. Summer 1986 and seasonal variation. Ciencias Marinas 17, 121–145.
80
Hernández-Carmona, G., Rodriguez-Montesinos, Y.E., Torres-Villegas, J.R., Sanchez-Rodriguez, I. & Vilchis,

M.A. 1989a. Evaluation of Macrocystis pyrifera (Phaeophyta, Laminariales) kelp beds in Baja Cali-
fornia, México. I. Winter 1985–1986. Ciencias Marinas 15, 1–27.
Hernández-Carmona, G., Rodriguez-Montesinos, Y.E., Torres-Villegas, J.R., Sanchez-Rodriguez, I., Vilchis,
M.A. & Garcia-de la Rosa, O. 1989b. Evaluation of Macrocystis pyrifera (Phaeophyta, Laminariales)
kelp beds in Baja California, México. II. Spring 1986. Ciencias Marinas 15, 117–140.
Hobday, A.J. 2000a. Abundance and dispersal of drifting kelp Macrocystis pyrifera rafts in the Southern
California Bight. Marine Ecology Progress Series 195, 101–116.
Hobday, A.J. 2000b. Age of drifting Macrocystis pyrifera (L.) C. Agardh rafts in the Southern California
Bight. Journal of Experimental Marine Biology and Ecology 253, 97–114.
Hobday, A.J. 2000c. Persistence and transport of fauna on drifting kelp (Macrocystis pyrifera (L.) C. Agardh)
rafts in the Southern California Bight. Journal of Experimental Marine Biology and Ecology 253,
75–96.
Hobson, E.S. & Chess, J.R. 2001. Influence of trophic relations on form and behavior among fishes and benthic
invertebrates in some California marine communities. Environmental Biology of Fishes 60, 411–457.
Hoffmann, A.J., Avila, M. & Santelices, B. 1984. Interactions of nitrate and phosphate on the development
of microscopic stages of Lessonia nigrescens Bory (Phaeophyta). Journal of Experimental Marine
Biology and Ecology 78, 177–186.
Hoffmann, A.J. & Santelices, B. 1982. Effects of light intensity and nutrients on gametophytes and gameto-
genesis of Lessonia nigrescens Bory (Phaeophyta). Journal of Experimental Marine Biology and
Ecology 60, 77–89.
Hoffmann, A.J. & Santelices, B. 1991. Banks of algal microscopic forms: hypotheses on their functioning
and comparisons with seed banks. Marine Ecology Progress Series 79, 185–194.
Holbrook, S.J., Carr, M.H., Schmitt, R.J. & Coyer, J.A. 1990. Effect of giant kelp on local abundance of reef
fishes: the importance of ontogenetic resource requirements. Bulletin of Marine Science 47, 104–114.
Hooker, J.D. 1847. The Botany of the Antarctic Voyage of H.M. Discovery Ships Erebus and Terror. I. Flora
Antarctica. London: Reeve Brothers.
Howe, M.A. 1914. The marine algae of Perú. Memoirs of the Torrey Botanical Club 15, 1–185.
Huovinen, P.J., Oikari, A.O.J., Soimasuo, M.R. & Cherr, G.N. 2000. Impact of UV radiation on the early
development of giant kelp (Macrocystis pyrifera) gametophytes. Photochemistry and Photobiology
72, 308–313.
Hurd, C.L., Durante, K.M., Chia, F.S. & Harrison, P.J. 1994. Effect of bryozoan colonization on inorganic
nitrogen acquisition by the kelps Agarum fimbriatum and Macrocystis integrifolia. Marine Biology
121, 167–173.
Hurd, C.L., Stevens, C.L., Laval, B.E., Lawrence, G.A. & Harrison, P.J. 1997. Visualization of seawater flow
around morphologically distinct forms of the giant kelp Macrocystis integrifolia from wave-sheltered
and exposed sites. Limnology and Oceanography 42, 156–163.
Inglis, G. 1989. The colonization and degradation of stranded Macrocystis pyrifera (L.) C. Ag. by the
macrofauna of a New Zealand sandy beach. Journal of Experimental Marine Biology and Ecology
125, 203–218.
Isaac, W.E. 1937. Studies of South African seaweed vegetation. I. West coast from Lamberts Bay to the Cape
of Good Hope. Transactions of the Royal Society of South Africa 25, 115–151.
Jackson, G.A. 1977. Nutrients and production of giant kelp, Macrocystis pyrifera, southern California. Lim-
nology and Oceanography 22, 979–995.
Jackson, G.A. 1987. Modeling the growth and harvest yield of the giant kelp Macrocystis pyrifera. Marine
Biology 95, 611–624.
Jackson, G.A. 1998. Currents in the high drag environment of a coastal kelp stand off California. Continental
Shelf Research 17, 1913–1928.
Jackson, G.A. & Winant, C.D. 1983. Effect of a kelp forest on coastal currents. Continental Shelf Research
2, 75–80.
Jensen, J.R., Estes, J.E. & Tinney, L. 1980. Remote sensing techniques for kelp surveys. Photogrammetric
Engineering and Remote Sensing 46, 743–755.
81
Jones, L.G. 1971. The biology of giant kelp beds (Macrocystis) in California: Studies on selected small
herbivorous invertebrates inhabiting Macrocystis canopies and holdfasts in southern Californian kelp
beds. Nova Hedwigia 32, 343–367.
Juhl-Noodt, H. 1958. Beiträge zur Kenntnis der Perúanischen Meeresalgen. I. Kieler Meeresforschungen 14,
167–174.
Kaehler, S., Pakhomov, E.A. & McQuaid, C.D. 2000. Trophic structure of the marine food web at the Prince
Edward Islands (Southern Ocean) determined by delta13C and delta15N analysis. Marine Ecology
Progress Series 208, 13–20.
Kain, J.M. 1979. A view of the genus Laminaria. Oceanography and Marine Biology: An Annual Review 17,
101–161.
Kain, J.M. 1982. Morphology and growth of the giant kelp Macrocystis pyrifera in New Zealand and California.
Kenner, M.C. 1992. Population dynamics of the sea urchin Strongylocentrotus purpuratus in a central Cali-
fornia kelp forest: recruitment, mortality, growth and diet. Marine Biology 112, 107–118.
Kim, S.L. 1992. The role of drift kelp in the population ecology of a Diopatra ornata Moore (Polychaeta:
Onuphidae) ecotone. Journal of Experimental Marine Biology and Ecology 156, 253–272.
Kimura, R.S. & Foster, M.S. 1984. The effects of harvesting Macrocystis pyrifera on the algal assemblage in
a giant kelp forest. Hydrobiologia 116/117, 425–428.
Kinlan, B.P., Graham, M.H. & Erlandson, J.M. 2005. Late-quaternary changes in the size and shape of the
California Channel Islands: implications for marine subsidies to terrestrial communities. Proceedings
of the California Islands Symposium 6, 119–130.
Kinlan, B.P., Graham, M.H., Sala, E. & Dayton, P.K. 2003. Arrested development of giant kelp (Macrocystis
pyrifera, Phaeophyceae) embryonic sporophytes: a mechanism for delayed recruitment in perennial
kelps? Journal of Phycology 39, 47–57.
Kopczak, C.D., Zimmerman, R.C. & Kremer, J.N. 1991. Variation in nitrogen physiology and growth among
geographically isolated populations of the giant kelp, Macrocystis pyrifera (Phaeophyta). Journal of
Ladah, L.B. & Zertuche-Gonzalez, J.A. 2004. Giant kelp (Macrocystis pyrifera) survival in deep water
(25–40 m) during El Niño of 1997–1998 in Baja California, México. Botanica Marina 47, 367–372.
Ladah, L.B., Zertuche-Gonzalez, J.A. & Hernández-Carmona, G. 1999. Giant kelp (Macrocystis pyrifera,
Phaeophyceae) recruitment near its southern limit in Baja California after mass disappearance during
ENSO 1997–1998. Journal of Phycology 35, 1106–1112.
Lafferty, K.D. 2004. Fishing for lobsters indirectly increases epidemics in sea urchins. Ecological Applications
14, 1566–1573.
Laidre, K.L., Jameson, R.J. & DeMaster, D.P. 2001. An estimation of carrying capacity for sea otters along
the California coast. Marine Mammal Science 17, 294–309.
Lane, C.E., Mayes, C., Druehl, L.D. & Saunders, G.W. 2006. A multi-gene molecular investigation of the
kelp (Laminariales, Phaeophyceae) supports substantial taxonomic re-organization. Journal of Phy-
cology 42, 493–512.
Lavoie, D.R. 1985. Population dynamics and ecology of beach wrack macroinvertebrates of the central
California [U.S.A.] coast. Bulletin of the Southern California Academy of Sciences 84, 1–22.
Lawrence, J.M. 1975. On the relationships between marine plants and sea urchins. Oceanography and Marine
Biology: An Annual Review 13, 213–286.
Leet, W.S., Dewees, C.M., Klingbeil, R. & Johnson, E.J. (eds) 2001. California’s living marine resources: a
status report. State of California Resources Agency and Fish and Game, Sacramento, California.
Leighton, D.L. 1966. Studies of food preference in algivorous invertebrates of southern California kelp beds.
Pacific Science 20, 104–113.
Leonard, G.H. 1994. Effect of the bat star Asterina miniata (Brandt) on recruitment of the giant kelp
Macrocystis pyrifera C. Agardh. Journal of Experimental Marine Biology and Ecology 179, 81–98.
Lewis, R. & Neushul, M. 1994. Northern and Southern Hemisphere hybrids of Macrocystis (Phaeophyceae).
Lewis, R.J., Neushul, M. & Harger, B.W.W. 1986. Interspecific hybridization of the species of Macrocystis
in California. Aquaculture 57, 203–210.
82
Lindberg, D.R. 1991. Marine biotic interchange between the Northern and Southern Hemispheres. Paleo-
biology 17, 308–324.
Lleellish, M., Fernández, E. & Hooker, Y. 2001. Disturbancia del bosque submareal de Macrocystis pyrifera
durante El Niño 1997–1998 en la bahía de Pucusana. In Sustentabilidad de la Biosiversidad: Un
problema actual, bases científico-técnicas, teorizaciones y perspectivas, K. Alveal & T. Antezana (eds).
Concepción, Chile: Universidad de Concepción, 331–350.
Lobban, C.S. 1978a. The growth and death of the Macrocystis sporophyte (Phaeophyceae, Laminariales).
Phycologia 17, 196–212.
Lobban, C.S. 1978b. Growth of Macrocystis integrifolia in Barkley Sound, Vancouver Island, B.C. Canadian
Journal of Botany 56, 2707–2711.
Lüning, K. 1990. Seaweeds. Their Environment, Biogeography and Ecophysiology. New York: John Wiley &
Sons.
Lüning, K. & Neushul, M. 1978. Light and temperature demands for growth and reproduction of Laminarian
gametophytes in southern and central California. Marine Biology 45, 297–310.
Macaya, E.C., Boltaña, S., Hinojosa, I.A., Macchiavello J.E., Valdivia, N.E., Vásquez N.R., Buschmann A.H.,
Vásquez, J.A., Vega, J.M.A. & Thiel, M. 2005. Presence of sporophylls in floating kelp rafts of
Macrocystis spp. (Phaeophyceae) along the Chilean Pacific Coast. Journal of Phycology 41, 913–922.
Maier, I., Hertweck, C. & Boland, W. 2001. Stereochemical specificity of lamoxirene the sperm-releasing
pheromone in kelp (Laminariales, Phaeophyceae). Biological Bulletin (Woods Hole) 201, 121–125.
Maier, I., Müller, D.G., Gassmann, G., Boland, W. & Jaenicke, L. 1987. Sexual pheromones and related egg
secretions in Laminariales (Phaeophyta). Zeitschrift Naturforschung Section C Biosciences 42, 948–954.
Mann, K.H. 1973. Seaweeds: their productivity and strategy for growth. Science 182, 975–981.
Mattison, J.E., Trent, J.D., Shanks, A.L., Akin, T.B. & Pearse, J.S. 1977. Movement and feeding activity of
red sea urchins (Strongylocentrotus franciscanus) adjacent to a kelp forest. Marine Biology 39, 25–30.
McCleneghan, K. & Houk, J.L. 1985. The effects of canopy removal on holdfast growth in Macrocystis
pyrifera (Phaeophyta; Laminariales). California Fish and Game 71, 21–27.
McConnico, L. & Foster, M.S. 2005. Population biology of the intertidal kelp, Alaria marginata Postels and
Ruprecht: a non-fugitive annual. Journal of Experimental Marine Biology and Ecology 324, 61–75.
McLean, J.H. 1962. Sublittoral ecology of kelp beds of the open coast near Carmel, California. Biological
Bulletin (Woods Hole) 122, 95–114.
McPeak, R.H. 1981. Fruiting in several species of Laminariales from southern California. Proceedings of the
International Seaweed Symposium 8, 404–409.
Moe, R.L. & Silva, P.C. 1977. Antarctic marine flora: uniquely devoid of kelps. Science 196, 1206–1208.
Moore, L.B. 1943. Observations on the growth of Macrocystis in New Zealand, with a description of a free-
living form. Transactions of the Royal Society of New Zealand 72, 333–340.
Moreno C.A. & Jara H.F. 1984. Ecological studies of fish fauna associated with Macrocystis pyrifera belts in
the south of Feuguian Islands, Chile. Marine Ecology Progress Series 15, 99–107.
Moreno, C.A. & Sutherland, J.P. 1982. Physical and biological processes in a Macrocystis pyrifera community
near Valdivia, Chile. Oecologia 55, 1–6.
Müller, D.G., Maier, I. & Müller H. 1987. Flagellum autofluorescence and photoaccumulation in heterokont
algae. Photochemistry and Photobiology 46, 1003–1008.
Muñoz, V., Hernandez-Gonzalez, M.C., Buschmann, A.H., Graham, M.H. & Vásquez, J.A. 2004. Variability
in per capita oogonia and sporophyte production from giant kelp gametophytes (Macrocystis pyrifera,
Phaeophyceae). Revista Chilena de Historia Natural 77, 639–647.
Neushul, M. 1959. Studies on the growth and reproduction of the giant kelp Macrocystis. Ph.D. dissertation,
University of California, Los Angeles, California.
Neushul, M. 1963. Studies on the giant kelp Macrocystis. II. Reproduction. American Journal of Botany 50,
354–359.
Neushul, M. 1971. The biology of giant kelp beds (Macrocystis) in California: the species of Macrocystis.
Nova Hedwigia 32, 211–222.
Neushul, M. & Haxo, F.T. 1963. Studies on the giant kelp, Macrocystis. I. Growth of young plants. American
Journal of Botany 50, 349–353.
83
Nicholson, N.L. 1978. Evolution within Macrocystis: Northern and Southern Hemisphere taxa. Proceedings
of the International Symposium on Marine Biogeography and Evolution in the Southern Hemisphere
2, 433–441.
Nisbet, R.M. & Bence, J.R. 1989. Alternative dynamic regimes for canopy-forming kelp: a variant on density-
vague population regulation. American Naturalist 134, 377–408.
North, W.J. 1971. The biology of giant kelp beds (Macrocystis) in California: introduction and background.
North, W.J. 1987. Biology of the Macrocystis resource in North America. In Case Studies of Seven Commercial
Seaweed Resources, M.S. Doty et al. (eds). San Francisco, California: FAO.
North, W.J. 1994. Review of Macrocystis biology. In Biology of Economic Algae, I. Akatsuka (ed.). Hague:
Academic Publishing, 447–527.
North, W.J. & Hubbs, C.L. (eds) 1968. Utilization of kelp-bed resources in southern California. State of
California Resources Agency and Fish and Game, Sacramento, California.
North, W.J., Jackson, G.A. & Manley S.L. 1986. Macrocystis and its environment: knowns and unknowns.
Aquatic Botany 26, 9–26.
North, W.J., James, D.E. & Jones, L.G. 1993. History of kelp beds (Macrocystis) in Orange and San Diego
Counties, California. Hydrobiologia 260/261, 277–283.
North, W.J. & Schaeffer, M.B. (eds) 1964. An investigation of the effects of discharged wastes on kelp.
Resource Agency of California, State Water Quality Control Board, Publication 26.
Norton, T.A. 1992. Dispersal by macroalgae. British Phycological Journal 27, 293–301.
Nuñez, L. & Vásquez, J.A. 1987. Amplitud trófica y utilización de microhábitat de 4 especies de peces
asociados a un bosque submareal de Lessonia trabeculata Villouta Santelices. Estudios Oceanológicos
(Chile) 6, 78–85.
Nyman, M.A., Brown, M.T., Neushul, M., Harger, B.W.W. & Keogh, J.A. 1993. Mass distribution in the
fronds of Macrocystis pyrifera from New Zealand and California. Hydrobiologia 260/261, 57–65.
Ojeda, F.P. & Santelices, B. 1984. Ecological dominance of Lessonia nigrescens (Phaeophyta) in central Chile.
Orr, M., Zimmer, M., Jelinski, D.E. & Mews, M. 2005. Wrack deposition on different beach types: spatial
and temporal variation in the pattern of subsidy. Ecology 86, 1496–1507.
Paddack, M.J. & Estes, J.A. 2000. Kelp forest fish populations in marine reserves and adjacent exploited areas
of central California. Ecological Applications 10, 855–870.
Papenfuss, G.F. 1942. Studies of South African Phaeophyceae. I. Ecklonia maxima, Laminaria, pallida,
Macrocystis pyrifera. American Journal of Botany 29, 15–24.
Papenfuss, G.F. 1964. Catalogue and bibliography of Antarctic and sub-Antarctic benthic marine algae. In
Biology of the Antarctic Seas, M.O. Lee (ed.). Washington, D.C.: American Geophysical Union, 1–70.
Pearse, J.S. & Hines, A.H. 1976. Kelp forest ecology of the central California coast. University of California,
Sea Grant College Program Annual Report 1975–76: Sea Grant Publication 57, 56–58.
Pearse, J.S. & Hines, A.H. 1979. Expansion of a central California kelp forest following the mass mortality
of sea urchins. Marine Biology 51, 83–91.
Pearse, J.S. & Hines, A.H. 1987. Long-term population dynamics of sea urchins in a central California kelp
forest: rare recruitment and rapid decline. Marine Ecology Progress Series 39, 275–283.
Perissinotto, R. & McQuaid, C.D. 1992. Deep occurrence of the giant kelp Macrocystis laevis in the Southern
Ocean. Marine Ecology Progress Series 81, 89–95.
Pfister, C.A. 1992. Costs of reproduction in an intertidal kelp: patterns of allocation and life history conse-
quences. Ecology 73, 1586–1596.
Powell, H.T. 1981. The ecology of Macrocystis and other kelps around the Falkland Islands (South Atlantic).
Proceedings of the International Seaweed Symposium 8, A48 only.
Raimondi, P.T., Reed, D.C., Gaylord, B. & Washburn, L. 2004. Effects of self-fertilization in the giant kelp,
Macrocystis pyrifera. Ecology 85, 3267–3276.
Reed, D.C. 1987. Factors affecting sporophyll production in the giant kelp Macrocystis pyrifera. Journal of
Experimental Marine Biology and Ecology 113, 60–69.
Reed, D.C. 1990. The effects of variable settlement and early recruitment on patterns of kelp recruitment.
Ecology 71, 776–787.
84
Reed, D.C., Amsler, C.D. & Ebeling, A.W. 1992. Dispersal in kelps: factors affecting spore swimming and
competency. Ecology 73, 1577–1585.
Reed, D.C., Anderson, T.W., Ebeling, A.W. & Anghera, M. 1997. The role of reproductive synchrony in the
colonization potential of kelp. Ecology 78, 2443–2457.
Reed, D.C., Brzezinski, M.A., Coury, D.A., Graham, W.M. & Petty, R.L. 1999. Neutral lipids in macroalgal
spores and their role in swimming. Marine Biology 133, 737–744.
Reed, D.C., Ebeling, A.W., Anderson, T.W. & Anghera, M. 1996. Differential reproductive responses to
fluctuating resources in two seaweeds with different reproductive strategies. Ecology 77, 300–316.
Reed, D.C. & Foster, M.S. 1984. The effects of canopy shading on algal recruitment and growth of a giant
kelp (Macrocystis pyrifera) forest. Ecology 65, 937–948.
Reed, D.C., Kinlan, B.P., Raimondi, P.T., Washburn, L., Gaylord, B. & Drake, P.T. 2006. A metapopulation
perspective on patch dynamics and connectivity of giant kelp. In Marine Metapopulations, J.P. Kritzer
& P.F. Sale (eds). San Diego, California: Academic Press, 353–386.
Reed, D.C., Laur, D.R. & Ebeling, A.W. 1988. Variation in algal dispersal and recruitment: the importance
of episodic events. Ecological Monographs 58, 321–335.
Reed, D.C. & Lewis, R.J. 1994. Effects of an oil and gas-production effluent on the colonization potential of
giant kelp (Macrocystis pyrifera) zoospores. Marine Biology 119, 277–283.
Reed, D.C., Neushul, M. & Ebeling, A.W. 1991. Role of settlement density on gametophyte growth and
reproduction in the kelps Pterygophora californica and Macrocystis pyrifera (Phaeophyceae). Journal
of Phycology 27, 361–366.
Reed, D.C., Schroeter, S.C. & Raimondi, P.T. 2004. Spore supply and habitat availability as sources of
recruitment limitation in the giant kelp Macrocystis pyrifera. Journal of Phycology 40, 275–284.
Rigg, G.B. 1913. The distribution of Macrocystis pyrifera along the American shore of the Strait of Juan de
Fuca. Torreya 13, 158–159.
Rodriguez, S.R. 2003. Consumption of drift kelp by intertidal populations of the sea urchin Tetrapygus niger
on the central Chilean coast: possible consequences at different ecological levels. Marine Ecology
Rosenthal, R.J., Clarke, W.D. & Dayton, P.K. 1974. Ecology and natural history of a stand of giant kelp,
Macrocystis pyrifera, off Del Mar, California. Fishery Bulletin 72, 670–684.
Rosenthal, Y. & Broccoli, A.J. 2004. In search of paleo-ENSO. Science 304, 219–221.
Roughgarden, J., Gaines, S. & Possingham, H. 1988. Recruitment dynamics in complex life cycles. Science
241, 1460–1466.
Sala, E. & Graham, M.H. 2002. Community-wide distribution of predator-prey interaction strength in kelp
forests. Proceedings of the National Academy of Sciences of the United States of America 99,
3678–3683.
Salinas, N. 2000. Macrocystis integrifolia (Laminariales: Phaeophyta) en el norte de Chile: distribución
espacio-temporal y fauna asociada. M.S. thesis, Universidad Católica del Norte, Coquimbo, Chile.
Santelices, B. 1990. Patterns of reproduction, dispersal and recruitment in seaweeds. Oceanography and
Marine Biology: An Annual Review 28, 177–276.
Santelices, B., Hoffman, A.J., Aedo, D., Bobadilla, M. & Otaiza, R. 1995. A bank of microscopic forms on
disturbed boulders and stones in tide pools. Marine Ecology Progress Series 129, 215–228.
Santelices, B. & Ojeda, F.P. 1984a. Effects of canopy removal on the understory algal community structure
of coastal forests of Macrocystis pyrifera from southern South America. Marine Ecology Progress
Series 14, 165–173.
Santelices, B. & Ojeda, F.P. 1984b. Population dynamics of coastal forests of Macrocystis pyrifera in Puerto
Toro, Isla Navarino, southern Chile. Marine Ecology Progress Series 14, 175–183.
Saunders, G.W. & Druehl, L.D. 1992. Nucleotide sequences of the small-subunit ribosomal RNA genes from
selected Laminariales (Phaeophyta): implications for kelp evolution. Journal of Phycology 28,
544–549.
Saunders, G.W. & Druehl, L.D. 1993. Revision of the kelp family Alariaceae and the taxonomic affinities of
Lessoniopsis Reinke (Laminariales, Phaeophyta). Hydrobiologia 260/261, 689–697.
Sauvageau, C. 1915. Sur la sexualité heterogamique d’une Laminaire (Saccorhiza bulbosa). Comptes Rendus
de l’Académie des Sciences Paris 161, 796–799.
85
Scagel, R.F. 1947. An investigation on marine plants near Hardy Bay, B.C. Provincial Department of Fisheries
Report 1, Victoria, British Columbia, Canada.
Schiel, D.R., Andrew, N.L. & Foster, M.S. 1995. The structure of subtidal algal and invertebrate assemblages
at the Chatham Islands, New Zealand. Marine Biology 123, 355–367.
Schiel, D.R. & Foster, M.S. 1986. The structure of subtidal algal stands in temperate waters. Oceanography
and Marine Biology: An Annual Review 24, 265–307.
Schiel, D.R. & Foster, M.S. 2006. The population biology of large brown algae: ecological consequences of
multiphase life histories in dynamic coastal environments. Annual Review of Ecology Evolution and
Systematics 37, 343–372.
Schiel, D.R., Steinbeck, J.R. and Foster, M.S. 2004. Ten years of induced ocean warming causes comprehensive
changes in marine benthic communities. Ecology 85, 1833–1839.
Schimmelmann, A. & Tegner, M.J. 1991. Historical oceanographic events reflected in carbon13 to carbon12
ratio of total organic carbon in laminated Santa Barbara [California, U.S.A.] Basin sediment. Global
Biogeochemical Cycles 5, 173–188.
Schmitz, K. & Lobban, C.S. 1976. A survey of translocation in Laminariales (Phaeophyta). Marine Biology
36, 207–216.
Schroeter, S.C., Dean, T.A., Thies, K. & Dixon, J.D. 1995. Effects of shading by adults on the growth of
blade-stage Macrocystis pyrifera (Phaeophyta) during and after the 1982–1984 El Niño. Journal of
Setchell, W.A. 1932. Macrocystis and its holdfasts. University of California Publications in Botany 16,
445–492.
Setchell, W.A. & Gardner, N.L. 1925. The marine algae of the Pacific Coast of North America. III. Melano-
phyceae. University of California Publications in Botany 8, 383–898.
Seymour, R.J., Tegner, M.J., Dayton, P.K. & Parnell, P.E. 1989. Storm wave induced mortality of giant kelp
Macrocystis pyrifera in southern California. Estuarine Coastal and Shelf Science 28, 277–292.
Shanks, A.L, Largier, J.L., Brink, L., Brubaker, J. & Hooff, R. 2000. Demonstration of the onshore transport
of larval invertebrates by the shoreward movement of an upwelling front. Limnology and Oceanog-
raphy 45, 230–236.
Skottsberg, C. 1941. Communities of marine algae in subantarctic and Antarctic waters. Kungliga Svenska
Vetenskapsakademien Handlingar 19, 1–95.
Smith, S.D.A. 2002. Kelp rafts in the Southern Ocean. Global Ecology and Biogeography 11, 67–69.
Soto, R. 1985. Efectos del fenómeno El Niño 1982–83 en ecosistemas de la I Región. Investigación Pesquera
(Chile) 32, 199–206.
Spalding, H., Foster, M.S. & Heine, J.N. 2003. Composition, distribution, and abundance of deep-water
(>30 m) macroalgae in central California. Journal of Phycology 39, 273–284.
Stebbins, T.D. 1986. Density, distribution, and feeding of the marine snail Norrisia norrisi (Mollusca: Gas-
tropoda) on the kelp Macrocystis pyrifera (Phaeophyta: Laminariales). Bulletin of the Southern
California Academy of Sciences 85, 69–73.
Stegenga, H., Bolton, J.J. & Anderson, R.J. 1997. Seaweeds of the South African west coast. Contributions
from the Bolus Herbarium, Number 18, 1–655.
Steneck, R.S., Graham, M.H., Bourque, B.J., Corbett, D., Erlandson, J.M., Estes, J. & Tegner, M.J. 2002.
Kelp forest ecosystems: biodiversity, stability, resilience and future. Environmental Conservation 29,
436–459.
Stephens, J.S., Jr., Larson, R.J. & Pondella, D.J., Jr. 2006. Rocky reefs and kelp beds. In The Ecology of
Marine Fishes: California and Adjacent Waters, L.G. Allen et al. (eds). Berkeley, California: Univer-
sity of California Press.
Stephens, J.S., Jr., Morris, P.M., Zerba, K. & Love, M. 1984. Factors affecting fish diversity on a temperature
reef: the fish assemblage of Palos Verdes Point [California, U.S.A.], 1974–1981. Environmental
Biology of Fishes 11, 259–275.
Swanson, A.K. & Druehl, L.D. 2000. Differential meiospore size and tolerance of ultraviolet light stress within
and among kelp species along a depth gradient. Marine Biology 136, 657–664.
Tegner, M.J. & Dayton, P.K. 1987. El Niño effects on southern California kelp forest communities. Advances
in Ecological Research 17, 243–279.
86
Tegner, M.J. & Dayton, P.K. 1991. Sea urchins, El Niños, and the long-term stability of southern California
kelp forest communities. Marine Ecology Progress Series 77, 49–63.
Tegner, M.J., Dayton, P.K., Edwards, P.B. & Riser, K.L. 1995a. Sea urchin cavitation of giant kelp (Macrocystis
pyrifera) holdfasts and its effects on kelp mortality. Journal of Experimental Marine Biology and
Ecology 191, 82–99.
Tegner, M.J., Dayton, P.K., Edwards, P.B. & Riser, K.L. 1996. Is there evidence for long-term climatic change
in southern California kelp forests? CalCofi Reports 37, 111–126.
Tegner, M.J., Dayton, P.K., Edwards, P.B. & Riser, K.L. 1997. Large-scale, low-frequency oceanographic
effects on kelp forest succession: a tale of two cohorts. Marine Ecology Progress Series 146, 117–134.
Tegner, M.J., Dayton, P.K., Edwards, P.B., Riser, K.L., Chadwick, D.B., Dean, T.A. & Deysher, L.E. 1995b.
Effects of a large sewage spill on a kelp forest community: catastrophe or disturbance? Marine
Environmental Research 40, 181–224.
Tegner, M.J. & Levin, L.A. 1982. Do sea urchins and abalones compete in California kelp forest communities?
In International Echinoderms Conference, A.A. Balkema (ed.). Rotterdam: Tampa Bay, 265–271.
Thiel, M. & Gutow, L. 2005a. The ecology of rafting in the marine environment. I: The floating substrata.
Oceanography and Marine Biology: An Annual Review 42, 181–264.
Thiel, M. & Gutow, L. 2005b. The ecology of rafting in the marine environment. II: The rafting organisms
and community. Oceanography and Marine Biology: An Annual Review 43, 279–418.
Thiel, M. & Vásquez, J.A. 2000. Are kelp holdfasts islands on the ocean floor? Indication for temporarily
closed aggregations of peracarid crustaceans. Hydrobiologia 440, 45–54.
Towle, D.W. & Pearse, J.S. 1973. Production of the giant kelp, Macrocystis, estimated by in situ incorporation
of 14C in polyethylene bags. Limnology and Oceanography 18, 155–159.
Tutschulte, T.C. & Connell, J.H. 1988. Feeding behavior and algal food of three species of abalones (Haliotis)
in southern California. Marine Ecology Progress Series 49, 57–64.
Utter, B.D. & Denny, M.W. 1996. Wave-induced forces on the giant kelp Macrocystis pyrifera (Agardh): field
test of a computational model. Journal of Experimental Biology 199, 2645–2654.
van Tüssenbroek, B.I. 1989a. Smooth-bladed Macrocystis (Laminariales, Phaeophyta) from the Falkland
Islands. Phycologia 28, 520–523.
van Tüssenbroek, B.I. 1989b. Observations on branched Macrocystis pyrifera (L.) C. Agardh (Laminariales,
Phaeophyta) in the Falkland Islands. Phycologia 28, 169–180.
van Tüssenbroek, B.I. 1989c. The life span and survival of fronds of Macrocystis pyrifera (Laminariales,
Phaeophyta) in the Falkland Islands. British Phycology Journal 24, 137–141.
van Tüssenbroek, B.I. 1989d. Seasonal growth and composition of fronds of Macrocystis pyrifera in the
Falkland Islands. Marine Biology 100, 419–430.
van Tüssenbroek, B.I. 1989e. Morphological variations of Macrocystis pyrifera in the Falkland Islands in
relation to environment and season. Marine Biology 102, 545–556.
van Tüssenbroek, B.I. 1993. Plant and frond dynamics of the giant kelp, Macrocystis pyrifera, forming a
fringing zone in the Falkland Islands. European Journal of Phycology 28, 161–165.
Vásquez, J.A. 1993. Effects on the animal community of dislodgement of holdfasts of Macrocystis pyrifera.
Pacific Science 47, 180–184.
Vásquez, J.A. & Buschmann, A.H. 1997. Herbivore-kelp interactions in Chilean subtidal communities: a
review. Revista Chilena de Historia Natural 70, 41–52.
Vásquez, J.A., Camus, P.A. & Ojeda, F.P. 1998. Diversidad, estructura y funcionamiento de ecosistemas
costeros rocosos del norte de Chile. Revista Chilena de Historia Natural 71, 479–499.
Vásquez, J.A., Castilla, J.C. & Santelices, B. 1984. Distributional patterns and diets of four species of sea
urchins in giant kelp forest (Macrocystis pyrifera) of Puerto Toro, Navarino Island, Chile. Marine
Ecology Progress Series 19, 55–63.
Vásquez, J.A, Vega, J.M.A. & Buschmann, A.H. 2006. Long term studies on El Niño-La Niña in northern
Chile: effects on the structure and organization of subtidal kelp assemblages. Journal of Applied
Vásquez, J.A., Véliz, D. & Pardo, L.M. 2001. Biodiversidad bajo las grandes algas. In Sustentabilidad de la
Biosiversidad: Un problema actual, bases científico-técnicas, teorizaciones y perspectivas, K. Alveal
& T. Antezana (eds). Concepción, Chile: Universidad de Concepción, 293–308.
87
Vega, J.M.A., Vásquez, J.A. & Buschmann, A.H. 2005. Population biology of the subtidal kelps Macrocystis
integrifolia and Lessonia trabeculata (Laminariales, Phaeophyceae) in an upwelling ecosystem of
northern Chile: interannual variability and El Niño 1997–1998. Revista Chilena de Historia Natural
78, 33–50.
Vetter, E.W. 1994. Hotspots of benthic production. Nature 372, 47–47.
Villegas, M.J. 2002. Utilización de hábitat por parte de Lontra felina (Molina, 1782) (Carnívora, Mustelidae)
en Isla Choros (Cuarta Región de Chile) en relación con la abundancia y distribución de presas.
M.S. thesis, Universidad Católica del Norte, Coquimbo, Chile.
Viviani, C.A. 1979. Ecogeografía del litoral Chileno. Studies on Neotropical Fauna and Environment 14,
65–123.
Watanabe, J.M. 1984a. Food preference, food quality and diets of three herbivorous gastropods (Trochidae:
Tegula) in a temperate kelp forest habitat. Oecologia 62, 47–52.
Watanabe, J.M. 1984b. The influence of recruitment, competition, and benthic predation on spatial distributions
of three species of kelp forest gastropods (Trochidae: Tegula). Ecology 65, 920–936.
Watanabe, J.M. & Harrold, C. 1991. Destructive grazing by sea urchins Strongylocentrotus spp. in a central
California kelp forest: potential roles of recruitment, depth, and predation. Marine Ecology Progress
Series 71, 125–141.
Westermeier, R. & Möller, P. 1990. Population dynamics of Macrocystis pyrifera (L.) C. Agardh in the rocky
intertidal of southern Chile. Botanica Marina 33, 363–367.
Wheeler, P.A. & North, W.J. 1981. Nitrogen supply, tissue composition and frond growth rates for Macrocystis
pyrifera off the coast of southern California. Marine Biology 64, 59–69.
Wheeler, W.N. & Druehl, L.D. 1986. Seasonal growth and productivity of Macrocystis integrifolia in British
Columbia, Canada. Marine Biology 90, 181–186.
Wing, B.L. & Clendenning, K.A. 1971. The biology of giant kelp beds (Macrocystis) in California: kelp
surfaces and associated invertebrates. Nova Hedwigia 32, 319–341.
Wing, S.R., Botsford, L.W., Ralston, S.V. & Largier, J.L. 1998. Meroplanktonic distribution and circulation
in a coastal retention zone of the northern California upwelling system. Limnology and Oceanography
43, 1710–1721.
Womersley, H.B.S. 1954. The species of Macrocystis with special reference to those on southern Australian
coasts. University of California Publications in Botany 27, 109–132.
Womersley, H.B.S. 1987. The Marine Benthic Flora of Southeastern Australia. II. Phaeophyta. Adelaide:
South Australian Government Printing Division.
Yoon, H.S., Lee, J.Y., Boo, S.M. & Bhattacharya, D. 2001. Phylogeny of Alariaceae, Laminariaceae, and
Lessoniaceae (Phaeophyceae) based on plastid-encoded RuBisCo spacer and nuclear-encoded ITS
sequence comparisons. Molecular Phylogenetics and Evolution 21, 231–243.
Zimmerman, R.C. & Kremer, J.N. 1986. In situ growth and chemical composition of the giant kelp, Macrocystis
pyrifera: response to temporal changes in ambient nutrient availability. Marine Ecology Progress
Series 27, 277–285.
Zimmerman, R.C. & Robertson, D.L. 1985. Effects of El Niño on local hydrography and growth of the giant
kelp, Macrocystis pyrifera, at Santa Catalina Island, California. Limnology and Oceanography 30,
1298–1302.
Zobell, C.E. 1971. The biology of giant kelp beds (Macrocystis) in California: drift seaweeds on San Diego
County beaches. Nova Hedwigia 32, 269–315.
88
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Taylor & Francis
HABITAT COUPLING BY MID-LATITUDE, SUBTIDAL,

MARINE MYSIDS: IMPORT-SUBSIDISED OMNIVORES
PETER A. JUMARS
School of Marine Sciences & Darling Marine Center, University of Maine,
193 Clark’s Cove Road, Walpole, Maine 04573, U.S.
E-mail: jumars@maine.edu
Abstract Mysids often dominate mobile benthic epifaunas of mid-latitude continental shelves.
Macquart-Moulin & Ribera Maycas (1995) reported that the six most abundant species on western
and southern European shelves are all strong diel migrators. Published daytime epibenthic sledge
(sled) data from the surf zone to the shelf edge matched with published behavioural data on the
most abundant species were used to test, confirm and extend that relationship to other coastal
regions and to identify an association of abundant migrators with species that are important in fish
diets. They also reveal another pattern: a correspondence between abundant surf-zone species and
species that dominate estuarine faunas seasonally. Population concentrations at estuary mouths,
sills of fjords and in the surf zone suggest a lifestyle dependent upon horizontal fluxes. Marine
mysids that migrate between habitats are chronically undersampled in the field, however, and are
underrepresented in food-web models. Unfortunately, no single methodology samples both pelagic
and benthic individuals well and nearly all shelf measurements so far reported must be considered
underestimates of local abundance. Mysids are major dietary components for many benthic and
pelagic fishes, mammals, cephalopods and decapods, often for key life stages, and often because
mysid migrations result in encounters with predators. Mysids can be extraordinarily omnivorous,
with demonstrated capabilities to digest cellulose and diets spanning macrophyte detritus, more
labile detritus, large microalgae, and smaller animals and heterotrophic protists. They can be
sufficiently abundant and active to play roles in sediment transport. Contributing factors to their
underappreciation have been the lack of fidelity of mysids to single habitats, coupled with higher
fidelity of investigators to the study of single habitats. Sampling with classical methods has been
problematic because of effective evasion by mysids, compounded by extreme patchiness associated
with mysid schooling. Their frequent absence from coastal and even estuarine food-web models
has not been more conspicuous because the combination of their migration and omnivory spreads
their feeding impacts and because they are subsidised by horizontally imported plankton and seston
and are themselves horizontally exported in the form of predator gut contents and biomass. They
clearly link pelagic and benthic food webs in two important and ecosystem-stabilising ways,
however, by feeding in both habitats and by succumbing in both habitats to both cruising and sit-
and-wait predators. Consideration of resource and predation gradients and limited data implicate
horizontal, diel migrations as well, extending these linkages, especially in the onshore–offshore
direction. Somewhat paradoxically, the same features that have made them difficult to study by
classical means, in particular schooling, diet breadth, ontogenetic change in diet and migration
between habitats, suit migrating mysids well to new, individual- or agent-based modelling
approaches. Moreover, benthic observatories deploying acoustic technologies with spatial and
temporal resolution sufficient to resolve individual migratory behaviours promise powerful tests of
such models.
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PETER A. JUMARS
Introduction
For nearly two centuries, observations of zooplankton vertical migrations have aroused curiosity
and elicited alternative and compound explanations (Pearre 2003). Selective forces evoking and
altering these migrations include vertical gradients in resources, in predation risks and in environ-
mental drivers of physiological rates (i.e., temperature and salinity). Such gradients in risks and
benefits can be even steeper within the bottom boundary layer, including its upper layers of sediment
(Boudreau & Jørgensen 2001), and laterally across fronts, than they are in the overlying water
column. The focus of this review is on migrations between benthic and pelagic habitats by a subset
of the animal community that may also move horizontally, both across and along isobaths, con-
necting more than two habitats. For that reason, in this review the more general term ‘habitat
coupling’ is used rather than benthic-pelagic coupling (Schindler & Scheuerell 2002).
Widespread use of echo sounders after the rapid advance of underwater acoustics in World War II
brought attention to the ubiquity of vertical migrations and specifically to the oceanic deep scattering
layer. Echo-sounder frequencies near 12 kHz that were useful for locating the bottom proved
sensitive to air bladders of fishes and siphonophores. Based partly on such observations, Vinogradov
(1962) developed a conceptual scheme subsequently dubbed ‘Vinogradov’s ladder’: although diel
migrations from deeper than 600 m are rare, many deeper-dwelling species migrate part of the way
to the surface, so that predatory interactions provide a chain or ladder for vertical redistribution of
energy and materials that daily extends to depths in excess of 1000 m in the open ocean. The
proliferation of acoustic Doppler current profilers (ADCPs, operating typically at 300–600 kHz;
Brierly et al. 1998) and of bioacoustic instruments designed to detect zooplankton at acoustic
frequencies typically ranging from 250 kHz to a few megahertz (e.g., Gal et al. 1999) is revealing
the ubiquity and intensity in shallow waters of an inherently more complicated phenomenon that
has been dubbed the shallow scattering layer (Kringel et al. 2003). In waters too shallow to hold
a deep scattering layer, animals from many taxa have evolved foraging patterns and morphologies
compatible with living in or on the bottom, usually during the day, and rising into the water column,
usually at night. Although there is no need for a vertical ladder where the water is a single rung
deep, early data already show the outlines of a horizontal or oblique, onshore–offshore ladder in
the coastal zone.
Though still quite limited in number, deliberate, multifrequency acoustic studies of shallow-
water migrators suggest that water-column abundances (depth-integrated biomasses) of these
migrants may frequently exceed those of the holoplankton. This suggestion led to a systematic
examination of corroborative evidence for the ecological importance of these migrants. For prag-
matic reasons, in this review analysis is limited to a single large taxon, the Mysidacea (commonly
known as opossum shrimp), that appears in shallow, mid-latitude seas and often dominates such
migrations. Similarly, the focus is limited to subtidal, coastal habitats of mid-latitudes and to species
that occur outside estuaries during at least some seasons of the year. Work in other marine, estuarine
and freshwater environments is cited selectively when comparable information was not at hand for
mid-latitude marine systems. Literature on freshwater species or (oligohaline and mesohaline)
estuarine endemics has not been reviewed for the simple reason that the importance of vertically
migrating mysids in these systems is widely appreciated (e.g., Rudstam et al. 1989, Kotta & Kotta
2001a, Viitasalo et al. 2001). To avoid inflation of inferred importance by selective extraction of
conspicuous examples of migration and to give some insight into migrations of individual species,
a two-step process was used. The first step was to identify a few regional studies of mysids notable
for the spatial or temporal extent (or both) of their epibenthic sledge sampling. Thus, this review
is also focused away from hard bottoms, caves and vegetation, all habitats well exploited by mysids
but ones requiring different census methods. The second step was to review characteristics of
migrations in the mysid species that dominated samples in these studies. In both steps emphasis
90
HABITAT COUPLING BY MID-LATITUDE, SUBTIDAL, MARINE MYSIDS
has been on studies published after the major review by Mauchline (1980), citing prior literature
primarily when a particular citation was omitted by Mauchline (1980) or when focusing on
information that was not summarised by Mauchline (1980).
Additional information was then reviewed, substantiating the importance of mysids to coastal
and estuarine ecosystems. Confluence of multiple lines of evidence for the importance of migrating
mysids to both benthic and pelagic systems proved compelling. They often dominate diets of both
pelagic and benthic fishes in coastal waters and estuaries, highlighting the multiple risks inherent
in the migratory lifestyle. Mysids also appear to be important to the population abundances of some
of their prey species, but for the most part mysids are remarkable dietary generalists when all life
stages and habitat phases are included, and so are underappreciated stabilisers of the communities
that they inhabit and transit (McCann & Hastings 1997). Recently, their migrations have been
implicated as an important factor in sediment dynamics (Roast et al. 2004). Major habitat changes
due to climate, introduced species or human intervention have often produced major changes in
mysid populations that resonate through the food web. Despite underlying differences between
mysid-containing food webs in fresh and marine waters, analogy with lake systems takes advantage
of their closed boundaries to assess effects of mysid introduction, which are detected both up and
down the food web. Another indicator of potential importance is the latitudinal range and habitat
diversity over which high abundances of even single species are found; Neomysis americana
(S.I. Smith, 1873) is abundant from Nova Scotia to Florida and from shelf habitats 100 m deep to
salt marshes; in the last century it was introduced to the Atlantic coast of South America, where
it has become an important food-web component. The importance of N. americana as food for
both benthic and pelagic fishes over a broad geographic range was recognised in its original species
description (Smith 1873).
The question naturally arises as to why, despite engaging, comprehensive treatments of their
capabilities and roles (e.g., Mauchline 1980) and intense and sustained interest among the specialists
cited in this review, mysids do not figure more prominently in fisheries and oceanographic models
and texts. The most direct comparison is with the largely holoplanktonic euphausiids, a group of
similar body size and also large dietary breadth (but less expansion into detritivory) as a group.
The summary by Mauchline (1980) of both groups followed a parallel structure for each. Tellingly,
his chapter on “Mysids in the marine economy” is half as long as its counterpart for euphausiids,
and only a small portion is devoted to shallow-water species that migrate. Biological oceanographic
textbooks in general give an even more lopsided treatment.
Reasons for this shortage of information are manifold. Shallow-water migrations are funda-
mentally more complicated than better-studied migrations in the open ocean or in coastal holo-
plankton because component populations in benthic and pelagic habitats cannot be studied by the
same means and often are not sampled by a single investigator. Their natural reference frame shifts
back and forth between an Eulerian fixed reference frame and a Lagrangian, water-mass-following
reference frame with the change between benthic and pelagic habitats, respectively, seriously
complicating description and analysis. Even when they stay within the pelagic or benthic habitat,
mysids are notoriously poorly captured because of their effective evasive behaviours. More subtly,
their lack of freely released eggs or larvae leaves no evidence of large mysid populations in low-
flow or small-aperture capture devices, such as continuous plankton recorders, that efficiently
recover those non- or weakly swimming life stages in euphausiids, decapods and fishes. Extreme
patchiness of mysid populations, reinforced by schooling behaviours, make precise abundance
estimates even more difficult to achieve than they are for non-schooling animals. The migratory
lifestyle gives mysids access to horizontally imported pelagic food sources and leads through
encounter to their local export as gut contents and assimilated biomass of fishes and decapods,
effectively camouflaging their importance to local food webs and energy budgets; a large net import
or export would be far more conspicuous.
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PETER A. JUMARS
Migrations between the sea bed and the water column also generate semantic difficulties. Mees
& Jones (1997) took a habitat point of view and defined the hyperbenthos as those animals living
in the water layer immediately above the bottom. In this sense, migratory mysids spend part of
their time as hyperbenthos. The term fails, however, to capture the range of habitats occupied by
migratory mysids because in clear, shallow waters without bottom cover in the form of crevices
or vegetation, mysids often bury themselves during daylight, disappearing from the hyperbenthos.
Mysid migrations also exhibit considerable plasticity, varying in timing, intensity and vertical extent
seasonally, night to night and with tides (e.g., Abello et al. 2005, Taylor et al. 2005). To pursue
these migrations further from a habitat perspective thus would require more elaborate terminology
than even the refinements proposed recently by Dauvin & Vallet (2006). Instead, in the present
review an alternative approach is adopted that may lead more readily to quantitative models and
predictions by taking the perspective of an individual migrating through habitats. It is noted that,
because mysids swim actively when pelagic and may do so at times during their benthic phases,
this perspective is not truly Lagrangian (in the normal physical oceanographic sense of tracking a
parcel of water), although it follows that same spirit of following the entity of interest. Seeking
the simplest terminology that has this behavioural focus, the term ‘emergence’ is used herein to
describe the overall vertical migration behaviour between habitats and more specifically the upward
component of the migration (leaving the distinction to context). This usage follows precedent for
those who have focused on the migratory behaviour rather than on community structure in the
hyperbenthic habitat (Saigusa 2001). When the shift is from pelagic to benthic, the term ‘re-entry’
is used in the current review, reflecting the author’s benthic background.
Two recent developments promise accelerated understanding of the role of migratory mysids.
One development is the continued evolution of bioacoustic instrumentation and its deployment
methods, particularly in the context of high-power, high-bandwidth ocean observatories. The second
advance is the rapid development of flexible, individual-based models (IBMs). Many of the same
features that have made mysids difficult to study make them excellent subjects for applications and
tests of IBMs (i.e., their schooling behaviours, their occupation of multiple habitats, their use of
multiple food resources and their shifts in behaviour during development) (Grimm & Railsback
2005, Grimm et al. 2005). The combination of new technologies and models promises accelerated
advances in understanding of the extents, causes and consequences of mysid migrations through
tests of predictions about habitat usage.
Migratory capabilities, schooling and their consequences

Credibility of evidence for migrations rests in some measure on the sensitivities of sensory mech-
anisms to guide them and on swimming capabilities. Mysids as a group are well endowed in both
of these categories (Mauchline 1980). The earliest (Carboniferous to Jurassic) mysids appear to
have been holopelagic, and the transition to emergence to have been marked by the evolution of
statocysts with mineralised statoliths (Ariani et al. 1993), likely associated with the fitness enhance-
ment of directional guidance in emergence and re-entry. Marine species generally (including Neomy-
sis americana) secrete fluorite (CaF2), whereas low-salinity estuarine and freshwater species gen-
erally secrete vaterite (a CaCO3 polymorph of calcite and aragonite), although particular species
provide exceptions to each generalisation that reflect their lineages (Ariani et al. 1993). Mysids
have major impact on the marine fluorine cycle (Wittman & Ariani 1996), and their statoliths may
be abundant enough in some fossil marine strata to warrant extraction (Voicu 1981). Likewise,
calcite (transformed vaterite) from statocysts represents a substantial fraction of some Miocene
Paratethys deposits in the Ponto-Caspian region, where use of calcium carbonate minerals appears
to have first evolved in mysids (Ariani et al. 1993). Emergent mysids thus appear to have been very
abundant in coastal ecosystems for a very long time, and they are still abundant enough to leave
92
detectable statoliths in modern shelf sediments (Enbysk & Linger 1966). In addition to a pair of
statocysts for vertical orientation, a less well-identified mechanism for sensing depth is present
(Rice 1961, 1964) that is sensitive to pressure changes equivalent to less than 1 m of water and
that probably enables observed tidal rhythms in activity cycles (Mauchline 1980, Saigusa 2001,
Gibson 2003, Taylor et al. 2005).
In terms of horizontal navigation, mysids have long been known to utilise polarised light
(Bainbridge & Waterman 1957, 1958), and movements of their stalked eyes are co-ordinated with
information from the statocysts (Neil 1975a,b,c). Contrary to opinion in many recent references,
polarisation (specifically e-vector orientation) is a useful indicator of solar azimuth throughout
continental shelf depths and through most of the day, with the highest information content near
dusk and dawn because of high inclination of the e-vector with respect to the horizontal (Waterman
2005). Seasonal onshore–offshore migrations have been inferred from asynchronous seasonal changes
in abundance across habitats (e.g., Bamber & Henderson 1994), and polarised light probably provides
the directional cue, although it often is not clear to what extent the asynchrony in local abundance
is due to migration versus seasonally changing, local differences in population growth and mortality
(Mees et al. 1993). For reasons that also are not yet clear, a majority of onshore–offshore migrators
show winter maxima offshore, extending in high abundance into shallower water and estuaries
during some or much of the period from spring to fall (Mauchline 1980). Diel homing to the same
location over smaller scales has been documented experimentally in reef mysids (Twining et al.
2000). Utilisation of estuarine circulations to help maintain horizontal position on intermediate
scales has also been observed (i.e., either an interaction of horizontal and vertical bias or directed
navigation) (e.g., Orsi 1986, Moffat & Jones 1993, Schlacher & Wooldridge 1994, Kimmerer et al.
1998a,b), although variation in such behaviours with local conditions from year to year can be
considerable (Kimmerer 2002), as can differences among mysid species at the same estuarine
location (Sutherland & Closs 2001). Retention-assisting, horizontal migrations also have been
observed during slack tides (Köpcke & Kausch 1996).
Many mysid species are documented to be strong swimmers. Sustained swimming at 10 body
lengths s−1 is not unusual, with bursts in some species exceeding 20 body lengths s−1 (Mauchline
1980). At these sustained speeds, diel vertical, diagonal or horizontal excursions on the order of
1 km would be feasible, depending on local flow velocities, so diel vertical migrations to the shelf
edge are well within mysid capabilities. Habitats with flow speeds in excess of sustainable swim-
ming speeds appear to be avoided, however, and mysids shelter behind flow obstructions and in
the most slowly moving water layer directly over the bottom (Roast et al. 1998, Lawrie et al. 1999).
Perhaps the most important point to emphasise in this introduction is the reason to focus on
both abundance and migration. A point forgotten all too easily is that ecological importance to
individuals of another species is usually a function of interspecific encounter rates (Hurlbert 1971),
themselves a product of areal or volumetric abundance times relative velocity (e.g., Jumars 1993).
The combination of good sensory guiding mechanisms and strong swimming capabilities would
tend toward ballistic encounter during organised migrations, an advantage in feeding but a disad-
vantage when being preyed upon (Visser & Kiørboe 2006).
Encounters in mysids are often modulated by schooling behaviours. Mysids use visual and
tactile senses to form and maintain both highly polarised schools and less polarised aggregations
or swarms (Ritz 1994). Very large aggregations of varying local density and orientation are termed
shoals (Clutter 1969). Typical mysid schools range from 1–10 m in linear dimensions and 1–15 m3
in volume (Ritz 1994). Near the bottom, school shapes often become planar, typically with more
than one layer of mysids and sometimes differing in vertical structure by sex and life stage. Moving
schools tend to be elongate, whereas stationary swarms (albeit containing milling individuals) are
more circular (when near the sea bed) or spherical (Clutter 1969, Wittman 1977, Ohtsuka et al.
1995). Schooling is typical of animals out from the cover of vegetation and swimming off the
93
PETER A. JUMARS
bottom (i.e., in the pelagic phase), even when only a few centimetres from the bottom, but schools
may maintain oriented, evenly spaced formation while on the bottom. Mysids on or near the bottom
typically orient into the current (Mauchline 1980). Densities in swarms are often near 105 of
individuals (ind.) m−3, with mean interindividual separation distances near 2 cm; for a single layer,
that spacing yields about 2500 ind. m−2 (Mauchline 1980). The first emergence event of the night
shows clear schooling and a constant ascent velocity dependent on depth and local light conditions,
but later emergence does not appear to be as organised; schooling may not be maintained through
the night (Kringel et al. 2003, Abello et al. 2005, Taylor et al. 2005). One function of schooling is
to reduce average risk per individual (Ritz 1994) to individual predators, although schooling
predators or large individual predators (e.g., whales) may be quite effective in the presence of mysid
schooling. It is clear from gut contents of benthic and pelagic fishes that migrating mysids still
incur fatal risk and that fitness loss must be counterbalanced by even greater gain from migrations
if the migrations persist.
Hence, the observations made by Macquart-Moulin & Ribera Maycas (1995) in an exhaustive
sampling programme of the pelagic phase of mysids throughout the water column in the north-
western Mediterranean take on particular significance: they observed that the most abundant mysids
found on the continental shelves of Europe are diel migrators between the sea bed and the pelagic
environment. Based on the integration of a large number of studies with varying types of sampling
gear over a long period, Macquart-Moulin & Ribera Maycas (1995) concluded that six species
showed high benthic abundance on the shelf: Gastrosaccus sanctus (Van Beneden, 1861), G. spinifer
(Goës, 1863), Anchialina agilis (G.O. Sars, 1877), Haplostylus lobatus (Nouvel, 1951), H. lobatus
var. armata (Nouvel, 1951), and H. normani (G.O. Sars, 1877). They provided compelling new
data from the region near Marseille of migration to the surface in all six of these taxa. Deprez et al.
(2005) regard Gastrosaccus sanctus as a synonym of G. spinifer and Haplostylus normani as a
synonym of H. lobatus. Macquart-Moulin & Ribera Maycas (1995) also found strong evidence of
offshore migration or transport of Anchialina agilis and Haplostylus lobatus, both captured over
bottoms 700–1000 m deep, where individuals are not known to occur on the bottom. They captured
pelagic Anchialina agilis in bathyal waters during the day and collected a high percentage of dead
animals, suggesting that occurrence in waters deeper than 500 m is an extension beyond suitable
habitat.
Methods of data collection

To identify recent published records of mysid abundance, three sources have been used in this
review: the Food and Agriculture Organisation of the United Nations’ Aquatic Sciences and
Fisheries Abstracts (ASFA), Thomson Scientific’s Web of Science and Google Scholar. The first
two sources are limited primarily to citations later than those in the review of Mauchline (1980),
but the third source is expanding rapidly into older literature. Into the search fields of the first two
databases, ‘mysi*’ was entered and a country name that has a continental shelf, or in the case of
the United States or Canada, a state or province name, respectively. For Google Scholar ‘mysid’
was used and the place name. For ASFA and the Web of Science, the ‘and’ is a Boolean operator.
For Google Scholar, it was omitted (as Google in general ignores small, common words unless
they are within explicit quotation marks). From the references returned, selected were those that
documented mysid abundance either over an extensive period (a year or more) or a broad geographic
area or both during daytime on the basis of epibenthic sledge samples. Many of these sledge studies
used multiple, vertically arrayed nets (e.g., Zouhiri et al. 1998) to get information on near-bottom
vertical distributions, biased to an unknown degree by species-specific escape responses. Such
samples are referred to as ‘vertically resolved, epibenthic-sledge samples’. From the data provided,
mysids have been ranked in terms of their abundances, selecting the one to five abundant and
94
frequent species, using a smaller number when a natural break point in abundance occurred (a
difference of an order of magnitude or more in absolute abundance), and using the largest number
when a long study over a large area showed consistent dominance of one species in at least one
location and season. In each case, the choices of taxa are explained. Species names in quotation
marks were then used as search terms in the same three databases to determine the migratory
behaviour of the most abundant species. In addition, species names were searched in the NeMys
database (Deprez et al. 2004, 2005) using the inclusive list of species names (valid and invalid) for
references on behaviour and as a further check for inclusivity of publications with extensive
sampling of field abundance and publications on migratory behaviour. The NeMys database was
also used as one source of taxonomic authority, indicated on first use of the species name in the
body of this review, and for some information (especially for European species) about geographic
and depth ranges. For brevity, depth ranges of the species are summarised only in tabular form
(Table 1). For consistency, only benthic capture records were used. Where taxonomic ambiguities
or disputes over synonymy might affect conclusions, all databases were searched under both names.
Table 1 Mysid species identified as abundant in epibenthic sledge samples, along with their
known depth ranges, diel migratory behaviours and the study that established their high abundance
Depth limits (m) Source of information
Species (respective citations) Diel migratory behaviour (citations) on abundance
Mesopodopsis 1–42 (Buhl-Jensen & Very strong swimmer, extends vertical Beyst et al. 2001
slabberi Fosså 1991, Beyst et al. distribution at night (Apel 1992, Wang &
2001) Dauvin 1994)
Schistomysis 1–116 (Buhl-Jensen & Moderately strong swimmer, extends vertical Beyst et al. 2001
spiritus Fosså 1991, Beyst et al. distribution at night (Apel 1992, Wang &
2001) Dauvin 1994)
Schistomysis 1–25 (Cornet et al. 1983, Weaker swimmer, extends vertical Beyst et al. 2001
kervillei Beyst et al. 2001) distribution at night (Apel 1992, Wang &
Dauvin 1994)
Anchialina agilis 2–493 (Bacescu 1941, Strongest swimmer and migrator to limits of Dauvin et al. 2000
Cartes & Sorbe 1995) its benthic depth distribution; most of the
population leaves the bottom every night
(Macquart-Moulin & Ribera Maycas 1995)
Gastrosaccus 1–260 (Lagardère & Strong swimmers and migrators (Macquart- Dauvin et al. 2000
spinifer Nouvel 1980, San Moulin & Ribera Maycas 1995)
Vicente & Munilla
2000)
Haplostylus 17–420 (Lagardère & Strong swimmers and migrators (Macquart- Dauvin et al. 2000
lobatus Nouvel 1980, Dauvin Moulin & Ribera Maycas 1995)
et al. 2000)
Haplostylus 10–150 (Lagardère & Strong swimmers and migrators (Macquart- Dauvin et al. 2000
normani Nouvel 1980, Dauvin Moulin & Ribera Maycas 1995)
et al. 2000)
Erythrops elegans 1–125 (Bacescu & May be a diel migrator (Vallet et al. 1995) Zouhiri et al. 1998
Schiecke 1974, Cunha
et al. 1997)
Schistomysis 6–407 (Brattegard & Collected in nighttime surface samples in some Zouhiri et al. 1998
ornata Meland 1997) seasons; may be a diel migrator (Sorbe 1991)
Leptomysis 5–512 (Elizalde et al. Strong migrator (Mauchline 1980, Kaarvedt Cornet et al. 1983,
gracilis 1991, San Vicente & 1989) Cunha et al. 1997
Munilla 2000)
(continued on next page)
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PETER A. JUMARS
Table 1 (continued) Mysid species identified as abundant in epibenthic sledge samples, along
with their known depth ranges, diel migratory behaviours and the study that established their
high abundance
Depth limits (m) Source of information
Species (respective citations) Diel migratory behaviour (citations) on abundance
Mysideis parva 120–519 (Bacescu & Non-migrator (Elizalde et al. 1991). No Cornet et al. 1983,
Schiecke 1974, Elizalde description found. Cunha et al. 1997
et al. 1991)
Neomysis 1–232 (Wigley & Burns Strong diel, tidally modulated migrator, but Wigley & Burns 1971
americana 1971) perhaps not to the full extent of its depth
range (Herman 1963, Brown et al. 2005,
Taylor et al. 2005)
Americamysis 4–179 (Wigley & Burns Strong diel migrator (Williams 1972) Wigley & Burns 1971
bigelowi 1971, Allen 1984)
Erythrops 16–450 (Petryashev Migrates at least in some environments Wigley & Burns 1971
erythrophthalma 2002a) (Brunel 1979)
Metamysidopsis 1–14 (Clutter 1967) Slight upward shift of population mode at Clutter 1967
elongata night (Clutter 1969)
Neomysis 1–210 (Petryashev 2005) Strong diel migrator (Kringel et al. 2003) Clutter 1967
kadiakensis
Xenacanthomysis 1–104 (Petryashev 2005) Caught in mid-water trawls (Wing & Barr Kim & Oliver 1989
pseudomacropsis 1977)
Neomysis rayii 1–79 (Petryashev 2005) Caught in mid-water trawls (Wing & Barr Kim & Oliver 1989
1977)
Acanthomysis 1–104 (Petryashev 2005) Poorly known Kim & Oliver 1989
stelleri
Archaeomysis 0–2 (Petryashev 2005) Strong diel migrators (Takahashi & Takahashi &
kokuboi Kawaguchi 1997) Kawaguchi 1997
Archaeomysis 1–50 (Hanamura 1997) Strong diel migrators (Takahashi & Takahashi &
japonica Kawaguchi 1997) Kawaguchi 1997
Iiella ohshimai 1–5 (Takahashi & Strong diel migrators (Takahashi & Takahashi &
Kawaguchi 1995) Kawaguchi 1997) Kawaguchi 1997
Nipponomysis 1–5 (Yamamoto & Undescribed? Hanamura &
ornata Tominaga 2005) Matsuoka 2003,
Yamamoto &
Tominaga 2005
Given demonstrated mysid capabilities for social aggregation and movement, reported maximal
local abundances per unit of volume of water are not very informative regarding typical regional
abundances, and documentation of consistently high abundance over a long time or broad region
is a better indicator of consistent importance. This review therefore focused on a subset of those
references that provide abundance estimates from epibenthic sledge samples taken during the
benthic phase (i.e., when individuals are most susceptible to capture by a sledge). Drawbacks are
that these studies varied widely in the geometries of the net mouth openings used and that many
of these papers reported only numbers per unit of volume filtered (as determined by flow meter).
No attempt was made to express abundances per unit of volume or per unit of area when the original
author did not do so. For ease of comparison, however, all areal or volumetric abundance estimates
given per total area or volume of tow were converted to numbers per square or cubic metre.
96
Regionally abundant mysids and their migration habits

European shelves
One of the most challenging environments to sample with respect to abundance and emergence
behaviours is the shallow subtidal and in particular the surf zone. In 15 monthly samples with a
bottom sledge hauled by hand from four sites in the Belgian surf zone, Beyst et al. (2001) found
average animal densities to exceed 15 ind. m−2 and to vary in ash-free dry weight (AFDW) from
3 to >30 mg m−2. Three quarters of individuals overall were mysids, primarily of three dominant
species (Mesopodopsis slabberi (Van Beneden, 1861), Schistomysis spiritus (Norman, 1860) and
S. kervillei (G.O. Sars, 1885)), and mysids dominated AFDW in some seasons. As is typical of
such estimates, sampling efficiency is unknown for this sledge with these species and is assumed
to be 100% for purposes of the calculation, so true densities must be higher. The three-species
group also dominates the Voor delta, where Gastrosaccus spinifer is also abundant (Mees et al.
1993).
Patterns of diel migration in Mesopodopsis slabberi are not as well known as might be expected.
The species was described from in situ observations (Wittman 1977) as active and colourless during
the day, showing no visible substrata preferences and changing leadership within schools sponta-
neously. Wittman (1977) also noted that schools did not appear to return to the same location and
that predator-evading swarms veered horizontally without changing depth unless the predator
attacked from above, a behaviour that should aid in capture by an epibenthic sledge. Daytime
schools swam up to 50 cm above the substratum. Although Wittman (1977) did not specifically
name M. slabberi in that context, he implied that nocturnal expansion among the mysids he studied
was the norm. Wang & Dauvin (1994) found M. slabberi in epibenthic sledge samples both night
and day and concluded from its upward skewed distribution among vertically resolved samples that
it is an active swimmer, consistent with the observations of Wittman (1977). Wang & Dauvin (1994)
caught more individuals in nighttime sledge samples but remarked that it might have been because
of increased capture efficiency (less evasion in the dark). Zouhiri et al. (1998) in another series of
epibenthic sledge samples found crepuscular peaks in capture of M. slabberi, consistent with the
idea of distribution broadening above the bottom at night (and perhaps net evasion in the light).
An inference consistent with most observations and directly supported by paired benthic and
pelagic samples in the Jade estuary is that M. slabberi is concentrated near the bottom during the
day but spreads into the water column at night (Apel 1992). This spreading includes a horizontal
component, into the intertidal zone of at least one estuary during the night (Colman & Segrove
1955). It is worth noting that whether a seaward expansion also occurs in surface waters is unknown.
In a long-term study of the polyhaline zone of the very turbid Gironde estuary, however, M. slabberi
was captured abundantly in surface waters during daylight (Castel 1993, David et al. 2005). In
addition, in the region of South African surf-zone diatom blooms M. wooldridgei Wittman, 1992
(closely enough related that it was previously identified as M. slabberi) also migrated onshore at
night to take advantage of sinking surf-zone diatoms carried offshore in rip currents (Webb &
Wooldridge 1990) at the same time that another mysid species, Gastrosaccus psammodytes Tatter-
sall, 1958, migrated offshore from its inner surf-zone, daytime habitat to take advantage of that
same resource (Webb et al. 1988). Mesopodopsis slabberi appears to migrate offshore in winter
but to include vertical migrations in its repertoire there at 20 m water depth (van der Baan &
Holthuis 1971). Even in winter, however, this species is observed inside but near the mouths of
some estuaries (Mees & Hamerlynck 1992), so the entire population does not migrate offshore
seasonally, and both migration and site-dependent mortality need to be examined as components
of the distributional shift. Seasonally, M. slabberi also enters tidal creeks of salt marshes at high
tides in sufficient abundance to be important as a prey species there (Hampel et al. 2003a), but it
97
PETER A. JUMARS
is unclear to what extent active behaviour versus passive advection is responsible (Hampel et al.
2003b). Recent molecular genetic work shows some genetic differentiation among populations in
the northeast Atlantic and Mediterranean and Black Seas (Remerie et al. 2006) and also shows
what may be an important mysid trait allowing rapid adaptation (i.e., high intrapopulation genetic
diversity).
Schistomysis spiritus and S. kervillei show similar migration patterns to Mesopodopsis slabberi,
including nocturnal vertical spreading (van der Baan & Holthuis 1971, Apel 1992, Wang & Dauvin
1994). Of these two congeners, nighttime expansion into very shallow water has been reported for
Schistomysis spiritus (Colman & Segrove 1955). Wang & Dauvin (1994) documented near-bottom,
daytime vertical distributions and near-bottom, nighttime spreading patterns in vertically resolved
sledge samples that allowed them to rank swimming activity as Mesopodopsis slabberi > Schisto-
mysis spiritus > S. kervillei, with Gastrosaccus spinifer in the same category as Schistomysis kervillei
and none of these mysids in their lowest activity category.
Mesopodopsis slabberi is the most widely distributed of the three species geographically,
ranging from Iceland in the Atlantic to North Africa, widely through the Baltic and Mediterranean
and into the Black Sea (Deprez et al. 2005). Both Schistomysis congeners have somewhat more
restricted geographic distributions than Mesopodopsis slabberi, with Schistomysis spiritus ranging
from the Baltic to northern France and S. kervillei ranging from the North Sea to the southern
Atlantic coast of France (with a report from northwest Africa), but its habitat distribution is
comparable, ranging from shallow estuarine to shelf depths (Deprez et al. 2005). Within the
North Sea, S. spiritus, S. kervillei and Mesopodopsis slabberi peaked in abundance near shore
(Dewicke et al. 2003). Late-summer abundances (all mysids combined) averaged near 30 m−3 and
30 mg AFDW m−3 in sledge samples from the nearshore zone. All three species, however, reached
even higher densities in the polyhaline and mesohaline zones of estuaries (e.g., Castel 1993, Wang
& Dauvin 1994, Delgado et al. 1997, Azeiteiro et al. 1999, Lock & Mees 1999, Dauvin et al. 2000,
Mouny et al. 2000, Wittman 2001, Drake et al. 2002, Dewicke et al. 2003). In terms of winter
distributions, all three species are known to occur inside estuaries (near the mouth of the Schelde;
cf. Mees & Hamerlynck 1992), in shallow coastal waters of warm regions (e.g., Lock & Mees
1999) and also offshore (van der Baan & Holthuis 1971).
In deeper waters of the English Channel and the European shelf, other mysid species become
dominant. Dauvin et al. (2000) presented a summary of 432 epibenthic sledge samples taken at
15 stations in the English Channel, including 3 stations within the Seine estuary, and covering the
years 1988–1996. Those three stations have been excluded from the analysis in this review, except
to note that they support the habitat pattern observed elsewhere for M. slabberi (i.e., shallow-water
marine plus polyhaline-mesohaline estuarine water). The indisputable dominant in terms of abun-
dance and frequency of occurrence outside the Seine estuary is Anchialina agilis, with mean
abundances >1 ind. m−3 at both of the deepest stations, a coarse sand off Roscoff and a medium
sand off Plymouth, both at 75 m depth. The species occurred at all the stations outside the Seine.
Four other species occurred at over one half of the non-estuarine stations and reached mean
abundances of at least 1 ind. m−3 at a minimum of one station: Gastrosaccus spinifer, Haplostylus
lobatus, H. normani and Schistomysis ornata (G.O. Sars, 1864). Other studies in the same region
by the same group of investigators appear consonant with these broad conclusions (e.g., Vallet et al.
1995, Vallet & Dauvin 1998, 2001), although Zouhiri et al. (1998) clearly showed Erythrops elegans
(G.O. Sars, 1863) to be co-dominant with Anchialina agilis and Schistomysis ornata in autumn
samples from the 75-m station near Plymouth, so Erythrops elegans has been added to the list of
species for investigation of migratory habits in this review. Samples off Arcachon, France (Cornet
et al. 1983), and off Aveiro, Portugal (Cunha et al. 1997), support the ubiquity and abundance of
Anchialina agilis at shelf depths ≤125 m and the inclusion of Erythrops elegans as a frequent and
abundantly caught mysid. They also support adding Leptomysis gracilis (G.O. Sars, 1864) as a
98
frequent co-dominant and occasional dominant at 52–125 m depth. Both of these sampling efforts
also found relatively high abundances of Mysideis parva Zimmer, 1915 at 85–120 m depth, so this
species has been added to the list of abundant species (Table 1).
Anchialina agilis, Gastrosaccus spinifer, Haplostylus lobatus and H. normani are included in
the list of six species provided by Macquart-Moulin & Ribera Maycas (1995), along with incon-
trovertible evidence of their emergence at night. As its specific name implies, Anchialina agilis is
an exceptionally strong swimmer. As for most shelf mysids, direct observations are rare. Wittman
(1977) described A. agilis during the day in diving depths to be inactive and colourless and to cling
to leaves of Zostera without showing any reaction to natural predators or to touch by a diver. This
description is incompatible with inferences from vertically resolved epibenthic sledge samples,
where Anchialina agilis is usually nearly uniformly distributed among vertically arrayed nets near
the bottom (e.g., Zouhiri & Dauvin 1996). Either the behaviour of this species varies spatially or
they show an eventual escape response to the sledge that is strong enough to randomise their vertical
distribution at the sledge mouth. Given their ubiquity, observations with a remote underwater vehicle
or other camera system should be feasible to resolve their daytime behaviours beyond comfortable
depths for divers. Both A. agilis and Haplostylus normani showed decreases of abundance in bottom
trawls (Zouhiri & Dauvin 1996) and increases in surface plankton tows (Macquart-Moulin & Ribera
Maycas 1995) at night, when Anchialina agilis showed remarkable concentration in the very surface
10 cm of the water column (Champalbert & Macquart-Moulin 1970). Few other species are so
reduced in abundance in nighttime epibenthic sledge samples (Zouhiri & Dauvin 1996), prompting
the conclusion that most of the A. agilis and Haplostylus normani populations undergo diel
migration (Macquart-Moulin & Ribera Maycas 1995, Vallet et al. 1995). Anchialina agilis has been
caught at the surface in water columns 1000 m deep, likely due to cross-isobath advection of surface
waters during emergence (Macquart-Moulin & Patriti 1993, Macquart-Moulin & Ribera Maycas
1995), yet it is difficult from extant data to exclude the possibility of an active horizontal component
to the migration as a contributor (Macquart-Moulin & Ribera Maycas 1995). Suggestive of acci-
dental expatriation is the capture in bathyal waters during the day of dead specimens (Macquart-
Moulin & Ribera Maycas 1995). Macquart-Moulin & Ribera Maycas (1995) also concluded that
the whole population of Gastrosaccus spinifer became pelagic at night.
Of the migrators discussed by Macquart-Moulin & Ribera Maycas (1995) and included under
the abundance criteria in this review, Anchialinja agilis is distributed from the North Sea to the
northern Mediterranean and is captured in estuaries only as stray specimens. Haplostylus lobatus
and H. normani, with synonymy that has been disputed (Deprez et al. 2005), together cover a range
from the Porcupine Bight to the northern Mediterranean and also are rare in estuaries. Gastrosaccus
spinifer ranges from Norway, in all the seas surrounding the British Isles and into the northern
Mediterranean and along the adjacent North African coast, with disjunct reports from Ivory Coast
and the South Shetland Islands in the Southern Ocean (Deprez et al. 2005). It is found further into
estuaries than the others but generally in reduced numbers compared with the nearby shelf (e.g.,
Buhl-Jensen & Fosså 1991).
Macquart-Moulin & Ribera Maycas (1995) did not list Erythrops elegans, Schistomysis ornata,
Leptomysis gracilis or Mysideis parva among the most abundant shelf mysids and thus did not
assess their migratory capabilities. Little published information is available on migration in Eryth-
rops elegans. Zouhiri et al. (1998) on the basis of vertically resolved epibenthic sledge samples
listed it as a weak swimmer, along with Gastrosaccus spinifer and Schistomysis ornata, because
it tended to be caught in the lower nets. Vallet et al. (1995), quoted in Zouhiri & Dauvin (1996),
suggested that Erythrops elegans migrates on a diel cycle (up at night) from the bottom boundary
layer to the surface.
Mauchline (1980) found negative evidence in the Clyde Sea and Loch Etive for diel migration
of S. ornata and in the Clyde for Erythrops elegans. He did not comment on migration of Mysideis
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PETER A. JUMARS
parva. In an extensive series of vertical plankton tows from the continental shelf of western France
over the span of 2 yr, Beaudouin (1979) reported Schistomysis ornata only from February tows
near the Gironde. Sorbe (1991) reported this species to be abundant from 91 to 179 m depth off
Arcachon in southwest France. Based on qualitative plankton tows over the 91-m station, he reported
vertical migration in this species and suggested based on size-frequency data that the species
migrates seasonally across isobaths. Zouhiri et al. (1998) found S. ornata and Erythrops elegans
in high abundance in the western English Channel, but their data showing nighttime disappearance
of mysids as a group from sledge samples (with crepuscular peaks in abundance) came from a
station in the eastern channel. They captured both Schistomysis ornata and Erythrops elegans
primarily in the lower two sampling nets of their epibenthic sledge, prompting the classification
of these two species in the group of weakest swimmers. Dauvin et al. (2000) captured Schistomysis
ornata only in nighttime sledge tows. Vallet & Dauvin (2001), however, captured roughly equal
biomasses of this species in day and night tows with peak autumn abundances reaching 21 ind. m–2.
Vallet et al. (1995), quoted in Zouhiri & Dauvin (1996), suggested that S. ornata remained plank-
tonic all day. Kaartvedt (1989) in the abstract of his study on mysid migration in Masfjorden,
Norway, listed S. ornata as a vertical migrator, but in that paper reported capture in 57 mostly
nighttime Isaacs-Kidd mid-water trawls of only 10 individuals (7 in the shallowest region sampled,
above a bottom 35–40 m deep and 3 singletons elsewhere) but made reference to unpublished data
from Kaartvedt et al. (1988) to support the conclusion of frequent migration. Earlier observations,
summarised by Tattersall (1938), suggested that a small subset of breeding individuals enters the
water column at night. Also supporting a case for reduced migration in this species compared with
the others already explored is its reported occurrence in the guts of relatively few fish species in
the three databases queried (i.e., Gibson & Ezzi 1980, Mauchline 1980, Astthorsson, 1985), but
Mauchline (1980) included several more predators, including herring (Clupea harengus), so the
issue of the degree of diel migration is not well settled in terms of the fraction of the population
participating, the seasonality of the phenomenon, its short-term frequency or the height above
bottom at which potentially enhanced swimming activity occurs. Schistomysis ornata also appears
to be able seasonally to congregate at a coastal front, presumably via horizontal migration, to take
advantage of the concentrated food resources there (Dewicke et al. 2002).
Mauchline (1980) cited abundant evidence of diel vertical migration in Leptomysis gracilis, a
highly mobile swarmer. Subsequent observations underscore the diel migratory activities of
L. gracilis (into the water column at night; e.g., Kaartvedt 1985, 1989). Zouhiri & Dauvin (1996),
however, captured more individuals in epibenthic sledges at night than during the day, suggesting
that part of the population stays on or returns to the bottom and that it may be more easily captured
at night. They also observed this species to be concentrated in the lower nets of vertically resolved
tows, nominally indicating lower activity but perhaps indicating a species-specific escape response.
Very little information is available on the migratory behaviour of Mysideis parva. Elizalde et al.
(1991) reported that the species was concentrated in the lower nets of their sledge samples,
indicating weak swimming ability, though nocturnal activity cycles have not been ruled out. In
terms of predator gut contents, it has been reported only from thornback rays (Raja clavata) (see
Mauchline 1980), lending some support to the conclusion that little migration occurs.
Erythrops elegans is somewhat narrowly distributed latitudinally in the North Atlantic (not
reported from northern Norway, Iceland or Morocco) but it is found broadly in the northern
Mediterranean. Although it is found in some fjords, it has not been reported from shallower estuaries
or from salinities much below that of sea water. Schistomysis ornata occurs in the North Atlantic
off Iceland, from Norway to France along the European west coast and in coastal seas surrounding
the British Isles. To the east it extends into the Baltic. It is also reported from Morocco, but not
from the Mediterranean. Like Leptomysis gracilis, it occurs frequently in fjords. Leptomysis gracilis
is distributed from Norway south around the British Isles, through the Baltic and broadly in the
100
northern Mediterranean. It occurs frequently in fjords and in some shallower estuaries. Mysideis
parva has been reported on only a few occasions and only at strictly marine sites stretching from
southern Ireland to the Ionian Sea.
Erythrops elegans, Schistomysis ornata and Leptomysis gracilis co-occur in western Norwegian
fjords, where their depth ranges are generally narrower and shallower than the inclusive ones quoted
in Table 1 (Fosså & Brattegard 1990). Erythrops elegans was collected by Fosså & Brattegard
(1990) at 32–100 m depth and had a median depth of occurrence of 40 m. In the fjords, Schistomysis
ornata and Leptomysis gracilis had depth ranges of 32–350 m and 32–166 m and median depths
of occurrence of 66 and 89 m, respectively. The four shallowest stations in this wide-area survey
were at 32, 40, 74 and 100 m depth. Even more interesting is the horizontal distribution in detailed
studies of a single fjord of western Sweden (Buhl-Jensen & Fosså 1991). Erythrops elegans,
Leptomysis gracilis, Mesopodopsis slabberi, Schistomysis ornata and S. spiritus all reached high
abundances on one or both of the sill stations. Erythrops elegans and Mesopodopsis slabberi
occurred only on the sill. Leptomysis gracilis had highest abundance on the sill but also occurred
throughout the fjord. Schistomysis ornata peaked on the shelf just beyond the sill (with second-
and third-ranked abundances on the sill) but was distributed across all stations except the shallowest,
most upstream station (depth 33 m). Erythrops erythrophthalma (Goës, 1864) showed a similar
pattern, with high abundance just outside the sill, at the inner sill station, and the highest abundance
shown by any mysid (11 ind. m−2) just inshore of the sill at 72 m depth.
Northwest Atlantic shelves

Regrettably, systematic, intensive epibenthic sledge sampling has not been practised so frequently
on other shelves. On the west side of the Atlantic, the collection of the U.S. National Marine Fisheries
Service (NMFS) from the U.S. Atlantic coast remains indisputably the most comprehensive (Wigley
& Burns 1971). It includes samples with an epibenthic sledge (‘bottom skimmer’) and 11 other
kinds of samplers. Although this study and a follow-up analysis (Wigley & Theroux 1981) included
abundant core samples from areas further south, sledge samples were limited to the region between
Nova Scotia and Long Island. The NMFS survey left no room for doubt about the single most
dominant species (Wigley & Burns 1971): “N[eomysis]. americana is the most common mysid
inhabiting the northeastern coastal waters of the United States and undoubtedly the most abundant
mysid in the western North Atlantic Ocean. …The NMFS [National Marine Fisheries Service]
collection originally contained over 2 million specimens…”. The next most abundant mysids in the
NMFS collection were Erythrops erythrophthalma, with 4573 specimens, and Americamysis bigelowi
(Tattersall, 1926), with 2031 specimens (Wigley & Burns 1971). No other species yielded >382
specimens. No areal or volumetric abundance estimates were attempted by Wigley & Burns (1971),
who were quite sensitive to issues of sampling bias (e.g., Wigley 1967).
Mauchline (1980) cited abundant evidence that Neomysis americana is a frequent migrator in
shallow water but noted (p. 74) that Whiteley (1948) “found no evidence of a regular diel migration
in Neomysis americana on Georges Bank where the depth at which they were living, as deep as
75 m, was greater than in the coastal regions where this species is known to migrate fairly regularly”.
Brown et al. (2005) from an extensive collection of zooplankton samples documented seasonal
emergence, peaking in April and May, over the period from 1995 to 1989. In most years, peak
abundances captured in these plankton samples were 0.1–1 ind. m−3. The present author and
coworkers have been collecting emergence-trap and acoustic data on this species in the Damariscotta
River estuary, in the U.S. mid-coast region of Maine for over 5 yr and at depth ranges of 10–20 m;
it undergoes diel, tidally modulated emergence from approximately late June until early November,
although emergence may be weak or absent on any particular day (Abello et al. 2005, Taylor et al.
2005, P.A. Jumars, unpublished observations). Neomysis americana appears to be a strongly diel
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PETER A. JUMARS
migrator in some seasons in most habitats from which it has been reported (e.g., Calliari et al.
2001). Neomysis americana also shows spectacular ability to aggregate (up to 2500 ind. m−2) in
bottom-water salinity fronts of estuaries (Schiariti et al. 2006).
Mauchline (1980) did not comment on migration in Americamysis bigelowi. The 10-year study
of Williams (1972) left no doubt, however, that A. bigelowi migrates above the bottom in large
numbers. Further support comes from its presence in the guts of Atlantic silversides (Menidia
menidia) in 20 m of water off New York (Warkentine & Rachlin 1989).
Mauchline (1980) cited Brunel (1979) for documentation of vertical migration of Erythrops
erythrophthalma in the Gulf of St. Lawrence. More recently, Carter & Dadswell (1983) reported
planktonic capture of E. erythrophthalma in the very turbid Saint John River estuary, New Brunswick,
year-round, including all life stages, but they took no benthic samples. Beaudouin (1979) reported
it from only three of her vertical plankton hauls off Gascogne during one winter. It is among the
species reported by Astthorsson (1985) from cod (Gadus morhua) guts, but the small number of
observations leaves the regularity and depth limits of its diel migratory status uncertain.
When this review was written, Deprez et al. (2005) had not entered many geographic data on
mid-latitude mysids outside Europe and Africa so, herein, original reports are cited instead. The
natural range of Neomysis americana is from the Gulf of St. Lawrence to northeastern Florida
(Williams et al. 1974). Such a broad geographic range that includes high abundances at both its
extremes of latitude strongly suggests a very successful opportunist or generalist, a conclusion
supported by its invasion of coastal waters and estuaries of South America, where it was first
reported from Uruguay (González 1974), but has spread south at least as far as San Blas, Argentina
(Orensanz et al. 2002). Some fishes have come to depend on this resource (e.g., Sardiña & Lopez
Cazorla 2005a,b), and some sympatric copepod populations have declined (Hoffmeyer 2004,
although she does not attribute the effect to N. americana; M.S. Hoffmeyer, personal communica-
tion). Americamysis bigelowi is known from Georges Bank southward to Florida (Wigley & Burns
1971). The species frequently co-occurs with, but in substantially lower abundance than, Neomysis
americana (e.g., Allen 1984). These two species are capable of substantial carnivory (Fulton 1982).
Americamysis bigelowi (as Mysidopsis bigelowi) was thought to range into the Gulf of Mexico to
the Texas coast, but is replaced by a pair of closely related species in the Gulf of Mexico (Price
et al. 1994): Americamysis alleni Price, Heard & Stuck, 1994 and A. stucki Price, Heard & Stuck, 1994.
The former species is found in poly- and mesohaline estuaries and the surf zone to 15 m, whereas the
latter species has a deeper distribution out to the shelf edge (Price et al. 1994). Deprez et al. (2005)
give distributional data for European Erythrops erythrophthalma, which is found off Greenland and
Svalbard, down the Norwegian coast and around the British Isles, off western France and along the
shelf and slope of the northern Mediterranean. The species is found southward along the U.S. east
coast as far as Delaware, peaking in abundance at 60–100 m depth in evidence from the NMFS
collection (Wigley & Burns 1971), and is widespread in the Arctic basin (Petryashev 2002a,b).
All three of the most abundant mysid species in the NMFS collection showed high abundance
on Georges Bank, with E. erythrophthalma most abundant on its southern flank, just above the
100-m isobath. Erythrops erythrophthalma is found primarily on the middle and outer shelf (Wigley
& Burns 1971, Petryashev 2002a). Both Neomysis americana and Americamysis bigelowi are
abundant in estuaries (e.g., Herman 1963, Allen 1984). In the NMFS survey, peak abundances on
the shelf for A. bigelowi were at 30–60 m depth (Wigley & Burns 1971). The bathymetric distri-
bution of Neomysis americana is unusual. In grab samples from the Gulf of Maine, Wigley and
Burns (1971) found this species at highest abundance from 30 to 60 m depth, noting its more
common presence in grabs taken during daylight. Within Cape Cod Bay, however, N. americana
apparently has a shallower abundance peak at 10–29 m depth (Maurer & Wigley 1982). In the
southern United States, the species rarely is captured in benthic samples offshore (Wigley & Burns
102
1971), occurring more frequently in the lower, middle and upper reaches of estuaries (e.g., Williams
1972, Zagursky & Feller 1985). This onshore shift, opposite in direction to that of many other
species with temperature or latitude, led Williams et al. (1974) to suspect (sub)speciation, but
morphological evidence did not support this interpretation, although the issue now merits re-
examination with molecular methods (Audzijonyte & Väinölä 2005, Remerie et al. 2006). Over
both the shelf and offshore shoals, within coastal embayments and near inlets, abundance peaks of
N. americana are reported in winter-spring, consistent with an overwintering generation breeding
then, although overwintering is not excluded as a possibility within estuaries as well (Carter &
Dadswell 1983). Estuarine abundance of N. americana generally peaks in summer from the mid-
Atlantic states northward, and the number of generations per year can increase to three in the
southern part of the species range (Cowles 1930, Whiteley 1948, Hulbert 1957, Herman 1963,
Hopkins 1965, Williams 1972). In South Carolina, N. americana is found year-round in subtidal
estuary channels and in shallow ocean waters, and its peak populations in estuaries are shifted
earlier in the year (DeLancey 1987, Johnson & Allen 2005, D.M. Allen, personal communication).
Northeast Pacific shelves

Elsewhere, information is much more fragmentary. In his classic study of nearshore mysids in the
surf zone of southern California, Clutter (1967) found the two most abundant mysids to be
Metamysidopsis elongata Holmes, 1900 (up to about 2000 ind. m−3 at 6 m below MLLW (mean
lower low water)) and Neomysis kadiakensis Ortmann, 1908 (up to about 180 ind. m−3 at 8 m below
MLLW); both are swarming species. Clutter (1969) reported only a subtle upward shift in median
population position of Metamysidopsis elongata at night, and he reported no observations on
Neomysis kadiakensis at night. Little more has been written after Clutter’s studies about the
behaviour and distribution of Metamysidopsis elongata except its essential fatty acid requirements
(Kreeger et al. 1991) and its role as a prey species for juvenile white seabass (Atractoscion nobilis)
(Donahoe 1997). A few observations on abundance patterns and laboratory culture of its Atlantic
subspecies have appeared (Tararam et al. 1996, Gama et al. 2002, and references therein).
Neomysis kadiakensis appears to follow an analogous pattern to the European Mesopodopsis
slabberi in abundance along the west coast of the United States. Clutter described Neomysis
kadiakensis as occurring in kelp beds as well as over open sand. This species can reach higher
abundances in estuaries. Dean et al. (2005) in a salt marsh within the San Francisco estuary noted
that it dominated mysid abundance there over the full year, with a spring peak in abundance at
244 ind. m−3. They measured a large net import of N. kadiakensis into the salt marsh, where
instantaneous mortality was calculated as 0.29 day−1. Kringel et al. (2003) in northern Puget Sound
over a muddy bottom at 20 m water depth observed coherent emergence and re-entry events of
N. kadiakensis associated with estimated biovolumes of 4–5 × 103 mm3 m−3. Moreover, these
nocturnal emergence events appear to have dominated the holoplankton in abundance (Kringel
et al. 2003). Vertical migration of this species is widespread in Puget Sound, but in the deeper
reaches individuals do not appear to migrate all the way to the sea bed (Thorne 1968). It seems
unlikely that Clutter (1969) could have missed such strong nocturnal emergence, so N. kadiakensis
likely differs in diel migration patterns along its range. Neomysis kadiakensis is distributed from
the Gulf of Alaska to southern California (Petryashev 2005).
Kim & Oliver (1989) specifically studied schooling crustaceans in regions where gray whales
(Eschrichtius robustus) fed in the Bering and Chukchi Seas. In diving observations concentrated
in the Bering Sea, they reported swarms of Xenacanthomysis pseudomacropsis Tattersall, 1933,
Neomysis rayii Murdoch, 1885 and Exacanthomysis arctopacifica Holmquist, 1981 and sampled
them with various means at depths from 3 to 24 m. Petryashev (1992) regarded E. arctopacifica
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PETER A. JUMARS
as a junior synonym of Acanthomysis stelleri Derzhavin, 1913. In Kim & Oliver’s study, Xenacan-
thomysis pseudomacropsis usually dominated and reached abundances of 600 ind. m−3. Acanth-
omysis stelleri was one to two orders of magnitude less abundant, and Neomysis rayii was reported
only from two sites at the southeast extreme of the Bering Sea, where it reached intermediate
abundances between those of the other two species. Direct observations of diel migration apparently
are lacking, but Wing & Bar (1977, quoted in Mauchline 1980) captured Xenacanthomysis
pseudomacropsis, Acanthomysis sp. and Neomysis rayii in mid-water trawls from the Chukchi Sea.
Neomysis rayii is a known winter diet component in common murres (Uria aalge) and marbled
murrelets (Brachyramphus marmoratus) off southeast Alaska (Sanger 1987, DeGange 1996) and
Acanthomysis spp. are listed as additional diet components of the latter (DeGange 1996), although
it is not clear how far the birds make excursions toward the bottom or mysids make excursions off
the bottom to effect their encounters. Neomysis rayii and Acanthomysis spp. are also taken by gray
whales (Eschrichtius robustus) further to the south, off Vancouver Island (Darling et al. 1998).
Neomysis rayii must have been caught often enough in plankton samples to be considered a pelagic
crustacean by some (McConnaughey & McRoy 1979), but simultaneous benthic and pelagic samples
over diel cycles would do much to clarify these issues for all three of the Alaskan species. Xena-
canthomysis pseudomacropsis occurs from central Japan to British Columbia, Neomysis rayii shares
that range and extends it to southern California, and Acanthomysis stelleri has the narrowest geo-
graphic range from northern Japan to the easternmost portion of the Aleutian peninsula. All three
species extend into the Chukchi Sea but not above the latitude of Wrangel Island (Petryashev 2002a).
Northwest Pacific shelves

Takahashi & Kawaguchi (1995, 1997) on the Pacific coast of northern Honshu, Japan, took isobath-
parallel, epibenthic sledge samples from the shallowest submerged station they could sample at the
lowest level of the spring tide out to 100 m from the tide line, to about 5 m water depth. Tows
were stratified by distance from shore in 10-m increments and were repeated monthly between
March 1992 and January 1993. Three species clearly dominated, with the dominant varying with
season and depth: Archaeomysis kokuboi Ii, 1964, A. japonica Hanamura, Jo & Murano, 1996 and
Iiella ohshimai (Ii, 1964). Archaeomysis kokuboi moved with the tide to stay in the shallowest
position, barely immersed, and showed a peak abundance of 511 ind. m−2 in 31 July in the 0- to
10-m range from the water’s edge. (The areal abundance estimate in the present review includes
no correction for sampling efficiency but simply divides the number in the tow by its 60-m2 area.)
Archaeomysis kokuboi emerged at night, expanding its distribution offshore, with the reproductively
most valuable members of the population showing less tendency to do so. Archaeomysis japonica
occupied the next depth stratum and did not migrate with the tides but also emerged at night. The
species reached peak abundance in the 10- to 20-m interval from the tide line in June at 60 ind. m–2.
Iiella ohshimai was found primarily in the deepest samples as juvenile stages but showed some
shoaling of its distribution in summer and also emerged at night. Iiella ohshimai reached peak
abundance of 1.3 ind. m−2 in the 20- to 30-m distance from the shoreline in August, but in other
months more than half of the captured individuals were found further offshore. All three species
spent daytime hours buried in the sand, and all three species showed depth segregation of life
stages. The Archaeomysis kokuboi population had breeding females as its shallowest members,
whereas the other two species’ populations had juveniles as their shallowest members. Archaeomysis
kokuboi and A. japonica are extensively exploited as food by surf-zone fishes and, as expected
from their burrowing behaviour during the day, are taken mostly at night by both benthic and
pelagic fishes that converge on this environment (Takahashi et al. 1999). Moreover, adult females
of A. kokuboi find spatial refuge in the extremes of shallow water (Takahashi et al. 2004).
104
Yamamoto & Tominaga (2005) took epibenthic sledge samples by boat from the shallow surf
zone of the Seto Inland Sea three times a month from May to August in three successive years.
Iiella ohshimai and Nipponomysis ornata Ii, 1964 co-dominated, with mean densities of 1.38 and
1.28 ind. m−2, respectively. One of the three dominant fish species fed primarily on the (daytime)
epibenthic species N. ornata, whereas the other two ingested Iiella ohshimai more frequently, based
on gut contents of daytime-collected fishes (Yamamoto & Tominaga 2005). In a separate study of
Japanese flounder (Paralichthys olivaceus) from the same region, Yamamoto et al. (2004) found
them to prefer the epifaunal Nipponomysis ornata. In another site in the same general region of
the Seto Inland Sea, sampled monthly by epibenthic sledge between May and October for two of the
same years, N. ornata showed much greater dominance over Iiella ohshimai, and mysid abundance
peaked at 250 ind. m−2 in May–June (Hanamura & Matsuoka 2003).
Of the four mysid species that dominated the nearshore subtidal in these Japanese studies, only
Archaeomysis kokuboi is listed by Petryashev (2005) in his biogeographic summary. He listed it
as being West Pacific, low boreal. Hanamura (1997) described its geographic limits as being from
central Hokkaido to Honshu in northern Japan and those of A. japonica as being shallow subtidal
to 50 m from Kyushu to Hokkaido. Suh et al. (1995) described an offshore migration in A. kukuboi
in eastern Korea in the afternoon, with onshore migration in the morning, but their study was done
before Hanamura (1997) resolved differences between some closely related species of Archaeomy-
sis. Hanamura et al. (1996) also confirmed the emergence of Archaeomysis japonica at night. It
appears that no information on diel migration of Nipponomysis ornata is available.
Other regions
In other regions, it was not possible to attempt the two-step methodology for lack of comparable
abundance estimates or information on migratory behaviour or lack of both, but it is clear that mysids
are ubiquitous at and beyond the range of latitudes considered here. Patagonian fjords, as but one
example, contain a rich mysid fauna (Brandt et al. 1997), with evidence of diel migrations (Antezana
1999), and merit more intensive study. In many cases, neuston or plankton samples confirm a pelagic
phase, but neither the connection to benthic populations nor phasing of migrations is known (e.g.,
Sawamto 1987). A few more of these geographically scattered reports are mentioned in the following
discussions of particular issues of ecological roles of mysids and drivers of their vertical migrations.
Mysid ‘umwelt’
Information in the three databases used is clearly biased geographically toward Europe and North
America, but where data are available on all three aspects, there is strong association among diel
migration by a mysid species, its high relative abundance among mysids in the same habitat and
its use as food by fishes and other animals. There is no reason to doubt that additional data would
add additional species to this list or that it could already be enlarged through other databases, but
some trends already are apparent. The clear risk during migration in species with this syndrome
must clearly be accompanied and outweighed by substantial fitness gains in nutrition, dispersal,
reproductive encounter and other aspects of life in order for these species to be among the most
abundant mysids. As but one example of these ‘other’ potential gains or reduced losses, mysids
living in macrophyte beds or over dense diatom mats may be driven out by low oxygen concen-
trations at night (Ledoyer 1969). One other pattern is apparent immediately from the data: many
of the abundant shelf species of mysids show even higher abundances in the convergence zones at
estuary mouths (Figure 1) than they do on shelves, with varying seasonal penetrations and popu-
lation irruptions into polyhaline, mesohaline and even oligohaline reaches of estuaries.
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PETER A. JUMARS
Erythrops elegans Americamysis alleni

Coastal
Erythrops erythrophthalma Mesopodopsis slabberi
Fjord plains estuary
Gastrosaccus spinifer Neomysis americana
with a sill with an inlet
Leptomysis gracilis Neomysis kadiakensis
Neomysis americana Schistomysis kervillei
Schistomysis ornata Schistomysis spiritus
Mesopodopsis
slabberi
Isobath
Americamysis bigelowi
Shelf edge
Increasing latitude
Figure 1 Species that meet abundance criteria in the text and that occur either in the surf zone and estuaries
or on the mid-shelf and in fjords or show both patterns (Neomysis americana). Mesopodopsis slabberi
apparently also invades fjords from the surf zone, whereas Americamysis bigelowi apparently invades estuaries
from the mid-shelf as surmised from seasonal abundance patterns.
Multiple drivers of emergence and their differing relative importance among species, locations
and times are as certain for mysids as they are for holoplankton (Pearre 2003), but strong involve-
ment of visual predators is clear from the characteristic pattern of emergence near dusk and re-entry
near dawn. What makes the emergent lifestyle unique is relative, Eulerian immobility during the
benthic phase, during which overlying waters are replaced. Holoplankton arguably can get a similar
subsidy in regions of high vertical shear, and euphausiids are routinely captured in epibenthic sledge
samples from mid-shelf depths and deeper (e.g., Cunha et al. 1997), but it will take a particular
combination of vertical shear and migration timing to remain in a region of high horizontal velocity
(e.g., Barber & Smith 1981). Daytime location of mysids under the surf zone, under inlets, on sills
(Figure 1) and under particular portions of estuaries (e.g., Kaartvedt 1989, Cunha et al. 1999) attests
to mysid virtuosity in utilising horizontal fluxes and resisting displacement. Horizontal subsidy
has been easiest to visualise over abrupt changes in topography such as seamounts (reviewed
by Genin 2004). The zooplankters that happen to occur over seamounts and shoals as their down-
ward diel migrations begin are subject to intense predation. In the mysid case, the topography can
be less steep, but the process is analogous and it is the mysids rather than their food taxa that do
the migrating; the large phytoplankton, other protists and small zooplankton that chance to be
advected above a mysid-rich area at night are subject to intense predation. Just as predators resident
on seamounts can produce patches of reduced zooplankton abundance in those waters that passed
over a seamount at night (Haury et al. 2000), nocturnal, pelagic patches of high mysid abundance
of diel migrators can be predicted to produce patches of reduced abundance in their holoplanktonic
prey. Preferred daytime mysid habitat underlies wave-driven currents, coastal currents, tidal cur-
rents, buoyancy-driven currents and topographically driven flow convergences over shoals, narrows
and sills, where resultant flows replace overlying waters at high frequency. The contrast between
mysids and euphausiids in fjords (Kaartvedt et al. 1988, Kaartvedt 1989) illuminates the benefits
of the mysid lifestyle. Euphausiids are clearly less capable of maintaining or returning to horizontal
106
co-ordinates of high horizontal flux or of high utility as refuge from predators and so are less able
to profit from them than are mysids.
Mysids thus fit into a broader pattern of horizontal import subsidy but have some special
adaptations that enhance their gains. Genin (2004, his ‘feed-rest’ hypothesis) noted that fishes on
seamounts can rest in the bottom boundary layer or behind flow obstructions when they are not
feeding. Mysids as omnivores can improve on feed-rest by feeding in the pelagic environment and
both feeding and resting on the bottom. They can cause ‘trophic focusing’ (Genin 2004) even where
topography is not steep. The reason that this focusing is not as evident as it could be, in turn, is
that mysids clearly suffer extensive mortality from mobile decapod and fish predators that hori-
zontally export much of the horizontal import subsidy that they gather. If mysids fed only on locally
produced prey and on a narrower range of resources, or if they were not fed on as heavily, their
impacts would be far easier to detect but they would be less important as stabilisers in the context
of new food-web theory (Montoya et al. 2006, Rooney et al. 2006). The likelihood that they return
to similar habitat but not the same location when they re-enter the benthic habitat suggests that
they also lend dynamic stability to benthic community structure; a local disturbance in terms of
mortality of mysids can recover literally overnight compared with the need in many other benthic
species for larval recruitment to occur.
Besides the typical pattern of daytime re-entry and nighttime emergence, a second indication
of the importance of visual predation as a driver of emergence is release from re-entry in especially
turbid waters (Carter & Dadswell 1983, Castel 1993) and association of high mysid abundances
with high turbidity zones of estuaries (e.g., Kimmerer et al. 1998b, Roast et al. 2004, Schiariti et al.
2006), which often leads to shoaling of populations in upper, more turbid reaches (e.g., Hulbert
1957). This association with and benefit from turbidity may underlie the paradoxical southward
shoaling of peak abundances of Neomysis americana on the continental shelf. South of Cape Cod,
particularly where barrier bars and islands develop along this passive continental margin, most fine
material delivered by rivers is trapped inside estuaries, and what little does get delivered has a short
residence time on the shelf. Dependence by juveniles on macrophyte detritus (either algal or
angiosperm) may also contribute to this southward shoaling because macroalgal substrata become
scarcer southward and plant and macroalgal fragments are among the particles largely trapped in
estuaries. Mysids are not absent from shallow or clear waters, but generally adopt one or more of
three strategies where they cannot hide within or below turbid waters: burying themselves in the
bottom, hiding in vegetation or other cover or schooling. If the reaction is to visual predation and
not turbidity per se, other optical phenomena that impede image formation will also benefit mysids
(e.g., image distortion through salinity or thermal microstructure) (suggested by M.J. Perry, personal
communication) and wave speckle and bubble clouds in the surf zone. Some mysids do congregate
at salinity fronts (Kotta & Kotta 2001b, Schiariti et al. 2006). Of course there are other potential
reasons, such as enhanced resource concentrations from electrostatically induced coagulation, from
salting out of organics or from frontal circulation patterns.
Day-night shifts in activity level (frequency and duration of movement) and height above the
bottom occur even in those species not known to migrate to surface waters, with increases typical
of nighttime (Fosså 1986). In the lowermost bottom boundary layer, because of the rapidly increas-
ing horizontal velocity and hence fluxes with increasing distance from the sea bed, even modest
changes in height above the bottom can yield large differences in exposure to resources. Coming
out of the sediments (for the mysid species that bury) as well as activity and height changes can
also produce major changes in encounter rates with predators. Saigusa (2001) provided a high-
resolution method for examining activity cycles, that is, by pump sampling from a depth 50 cm
below the water surface simultaneously with pump sampling 50 cm above the bottom and collecting
sequential 30-min samples from both streams. Saigusa (2001), at Akkeshi on the Pacific Coast of
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PETER A. JUMARS
Hokkaido in water ~3 m deep at high tide, Nipponomyis toriumi (Murano, 1977), in 25 days of
continuous sampling, showed a strong nocturnal periodicity in capture 0.5 m off the bottom, whereas
the surface pump showed weak or no diel periodicity. Both sampling streams showed tidal period-
icity in capture rates. At Ushimado on the Seto Inland Sea, in water of similar depth, where only
surface water was sampled, an 18-day time series showed Siriella japonica Ii, 1964 to have distinct
nocturnal periodicity in capture, again modulated by tides. A great deal of diversity in tidal and
diel periodicity and height of emergence above the bottom is expected of mysids across species,
locations and times as there is ample modulation of both risks and benefits on diverse scales.
Particularly striking in the time series analysis by Saigusa (2001) is the ubiquity of periodicity in
pump capture of potential prey of mysids and thus, in potential, for their capture by mysids.
An appreciation of mysids
Further evidence of mysid importance in the coastal marine economy
Food-web roles
Roles that mysids play in feeding their predators are increasingly recognised. A search on ‘mysi*’
and ‘feeding’ in ASFA currently returns about 900 citations of which >90% concern mysids in
diets of other animals. Mysids constitute particularly large fractions in the diets of many fishes in
the 3–15 cm length category. In a collection of nearly 500 beam trawl samples from the Wester-
schelde estuary (Hostens & Mees 1999), mysids occurred in >50% of the (mostly juvenile) fish
stomachs analysed and constituted >10% of the diets of two goby species (Pomatoschistus lozanoi
and P. minutus), garfish (Belone belone), two gadids (bib, Trisopterus luscus, and whiting, Mer-
langius merlangus), two flatfish species (Pleuronectes platessa and Platichthys flesus), herring
(Clupea harengus), seabass (Dicentrarchus labrax), sea snail (Liparis liparis), hook-nose (Agonus
cataphractus) and tub gurnard (Trigla lucerna). Beach seining in the southern Sea of Japan produced
similar dietary prominence in the 19 fish species recovered, with 67% of individuals feeding on
mysids (Inoue et al. 2003). As other prominent examples of mysid dominance in gut contents,
juvenile cod (Gadus morhua) <10 cm long in one study from the northwest Atlantic feed almost
exclusively on mysids (Link & Garrison 2002), and European hake (Merluccius merluccius) show
similar dietary preference (Papaconstantinou & Caragitsou 1987, Bozzano et al. 1997). Plummer
et al. (1983) examined gut contents of California halibut (Paralichthys californicus) trawled from
6–30 m depths off southern California and found Neomysis kadiakensis to dominate the diets of
fish <25 cm long. Patterns inside estuaries are similar (e.g., Nemerson & Able 2004). Particularly
when locally dominant mysid species bury themselves during the day and the water column is
shallow, bottom fishes may show higher feeding rates on mysids at night for simple encounter-rate
reasons (e.g., Hirota et al. 1990, Takahashi et al. 1999). Even mysids that stay near the bottom in
deeper water may become more active at night than during the day (Fosså 1986), increasing their
encounter rates with demersal fishes.
As an example of pelagic fish diets, mysids are important items in the nighttime diets of
European anchovies (Engraulis encrasicholus) (see Tudela & Palomera 1997), emphasising the
role of migration in encounter. In the Baltic, where the interaction of mysids and herring (Clupea
harengus) has been studied most extensively, herring feed on mysids primarily as medium to large
fish during winter (Möllmann et al. 2004). Whereas herring are widely regarded to be primarily
visual feeders and thus unlikely to eat nocturnally emerging mysids (Flinkman et al. 1992), other
clupeids do not necessarily follow this pattern. Alewives (Alosa pseudoharengus) use the lateral
line to locate mysid prey in the dark (Janssen et al. 1995) and in their sometimes-turbid marine
and estuarine habitats, specialise on mysids, crangonids and amphipods (Stone & Daborn 1987).
108
Moreover, gut contents analysis underestimates mysid contribution to assimilation because mysids
are more digestible than are many other taxa (Lankford & Targett 1997). The high dietary value
of mysids, based on richness in polyunsaturated fats (Navarro & Villanueva 2000, Richoux et al.
2004), has also been recognised in aquaculture feeds (e.g., Takeuchi et al. 2001). A particularly
clever application of stable carbon and nitrogen isotopic methods to flounders that eat mysids
permits estimation of their cumulative food consumption after release from a hatchery and docu-
mentation of their competition with natural stocks (Tominaga et al. 2003, Tanaka et al. 2005).
Another line of evidence for food-web significance comes from estuarine and coastal habitat
changes that have gone beyond their ‘normal’ limits through interannual variability (e.g., in fresh-
water runoff) or through habitat expansion of invasive species. A notable example is invasion of
the San Francisco Bay estuary system by the overbite clam (Potamocorbula amurensis), which
appears to have greatly reduced phytoplankton and small zooplankton stocks available to benthos
and plankton alike. Mysid stocks have plummeted to 10% of previous levels (Kimmerer & Orsi
1996), apparently as a result of food limitation (Orsi & Mecum 1986, Kimmerer 2002). Eight of
13 fish species showed declines in apparent response to this reduction, and only striped bass (Morone
saxatilis) contained more than traces of mysid remains (Feyrer et al. 2003) after the invasion.
Other vertebrates also utilise mysid prey. Notably, gray whales (Eschrichtius robustus) are
observed to feed on mysids both on shallow shelf feeding grounds (Kim & Oliver 1989) and along
the coast of British Columbia (Darling et al. 1998, Dunham & Duffus 2002, Stelle 2002, Patterson
2004). Bowhead whales (Balaena mysticetus) also take mysids (Lowry & Burns 1980). Seabirds
also utilise mysid populations (e.g., Moran & Fishelson 1971, Divoky 1978, Sanger 1987, DeGange
1996), and some species of polar seals (i.e., crabeater seals, Lobodon carcinophagus, and Weddell
seals, Leptonychotes weddellii) also have been found to eat mysids (Green & Williams 1986, Lake
et al. 2003). Seals at lower latitudes appear more commonly to have a less-direct dietary interaction
with mysids, experiencing parasitism by nematodes that use mysids and the fish predators of mysids
as intermediate hosts (Jackson et al. 1997). Humans also harvest mysids and do so on a commercial
scale (2000–3000 tons of one species per year) in Japan (Omori 1978).
Mysids are also substantial dietary components of many invertebrates. They appear critical, for
example, to diets of the smallest juvenile cuttlefish (Sepia officinalis) <2 cm long (Le Mao 1985)
and are substantial components in diets of many cephalopods (e.g., Aronson 1989, Huang 2004).
Many decapod shrimp species routinely prey on mysids, but among the most interesting couplings
is the frequent association of species of Crangon with vertically migrating mysids. Although it is
clear that Crangon spp. are not obligate feeders on mysids, where mysids are abundant local Crangon
species appear to specialise on them (e.g., Price 1962, Sitts & Knight 1979, Siegfried 1982, Pihl
& Rosenberg 1984, Wahle 1985, Hong & Oh 1989, Oh et al. 2001, Hanamura & Matsuoka 2003).
Crangon septemspinosa emerges on a diel cycle with or shortly after Neomysis americana according
to Taylor et al. (2005) and broadly overlaps its habitat and geographic ranges. On Georges Bank,
catches of the two species in macrozooplankton samples are highly correlated (Brown et al. 2005).
Taylor et al. (2005) pointed out that mysid emergence in their system is also associated with copepod
emergence. This copepod-mysid-crangonid emergence combination (Figure 2) greatly expands the
size spectrum of prey available in this movable feast, and some fish species use a substantial fraction
of the total range at one time or ontogenetically (e.g., Stone & Daborn 1987, Gatlin 1997, St.-Hilaire
et al. 2002, Yamamoto et al. 2004, Yamamoto & Tominaga 2005). Other invertebrates may also
follow the migration (e.g., Matsumoto 1995). It is also clear that the depth limits and sensory
postures of some benthic invertebrate predators are tuned to the arrival of vertical migrants (Lagar-
dère 1977). Mysids are both preyed upon and parasitised by protists (Buchanan & Hedley 1960,
Shields 1994). Mysids may also be eaten by other mysids of the same (Johnston & Ritz 2001,
Quirt & Lasenby 2002) or different species (Jerling & Wooldridge 1995). Mysid-mysid predation
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PETER A. JUMARS
Pelagic
Pelagic fishes
by night
Crangon
septemspinosa
Neomysis
americana
Mesozooplankton Emergent
meiofauna
Re-entry
Large
phytoplankton
Emergence
Benthic Demersal
by day fishes
Crangon
septemspinosa
Neomysis
americana
Other Emergent
meiofauna meiofauna
Microphytobenthos Detritus
Figure 2 The immediate food web of Neomysis americana, showing changes and some surprising constancy
over the diel cycle. The components within the dotted line all migrate, providing a much larger size spectrum
for encounter by larger decapods and fishes than would a single migrant. Mysids are extremely omnivorous
and are consumed by many species; the non-migrating part of this web is highly aggregated in the diagram.
is a function not only of size, but also of behaviours and abundances of the predator and prey
mysids as well as of alternative prey (Winkler & Greve 2004).
Because mysid diets have recently been reviewed by Takahashi (2004), present comments are
limited to a short summary. He pointed out that carnivory on holozooplankton is typical of later
juvenile and adult stages of mysids. In coastal and poly- to mesohaline estuarine environments,
prey are often copepods ingested at roughly 3–30% of mysid body carbon day−1. In oligohaline
and freshwater environments, cladocerans are often preyed upon heavily, probably because they
can be filtered in bulk (Viitasalo et al. 2001) and digested readily, and are ingested at 24–300% of
mysid body carbon day−1. The carnivorous component of the diet appears to be especially important
for growth and reproduction (Viherluoto et al. 2000). Smaller, younger juveniles that migrate eat
smaller holozooplankton, detritus and phytoplankton, but rarely ingest cells or chains <10 mm
long, often selecting diatoms in preference to other phytoplankton taxa. When large diatoms are
scarce, diets of juveniles may shift to microzooplankton (Jerling & Wooldridge 1995). The smallest
mysid individuals often do not migrate off the bottom and thereby may focus even more strongly
on a detrital or microphytobenthic diet. The potential dietary role of benthic diatoms for multiple
life stages of mysids bears emphasis (Mauchline 1968, Perissinotto et al. 2003). Takahashi (2004)
usefully described the trend of increasing carnivory with size and maturity as life-history omnivory,
supporting the generality of the stable isotope-based conclusions and terminology of Branstrator
(2000) for Mysis relicta Loven, 1862. The same term has been applied to fishes (Montoya et al.
2006), but mysid omnivory brackets a larger range of food types closer to the base of the food
web, yet spans more than one habitat. Mysid life-history omnivory is particularly well documented
by analysis of fatty acids (Richoux et al. 2005), and lipid accumulation in mysids from nighttime
predation on holozooplankton is probably enhanced by post-feeding migration to cooler waters
(Chess & Stanford 1999). Benthic phases may also span the entire gamut of diets from detritus to
microphytobenthos to heterotrophic protists and rotifers to larger meiofauna and even macrofauna
110
(Johannsson et al. 2001, Albertsson 2004), but all sizes of many species are more prone to ingest
detritus during the benthic phase than during the pelagic (Table 2 of Takahashi 2004). It seems
very likely that benthic-phase mysids are able to take advantage of the much greater food value of
suspended detritus (Mayer et al. 1993) either by direct suspension feeding from material already
suspended in the bottom boundary layer or by resuspending it from the bottom and then filtering
it (Mauchline 1980, Viherluoto et al. 2000).
Cellulases that are apparently endogenous have been reported in Mysis stenolepis S.I. Smith,
1873 by Friesen et al. (1986), and macrophyte detritus appears to be particularly important in the
winter diet of Neomysis americana in salt marshes in the southern part of its range (Zagursky &
Feller 1985). Stable isotopic data also support a winter shift toward a greater contribution from
refractory terrestrial, angiosperm detritus to mysid diets in coastal British Columbia (Mulkins et al.
2002). Froneman (2001), however, found no seasonality in the isotopic signature of Mesopodopsis
wooldridgei Wittman, 1992 in the temperate Kareiega estuary of South Africa but did find evidence
of likely contribution to its nutrition from eelgrass. All these studies contrast with the stable isotopic
results of Dauby (1995) for four species of Leptomysis that, contrary to the diel migrators focused
on in this review, school near the bottom during the day, remaining largely immobile relative to
the bottom and not feeding, and migrate down to feed on detritus at night; he found only minor
dietary influence from seagrass, with much greater reliance on micro- and macroalgae. Dauby’s
results, in turn, are compatible with those of Metillo & Ritz (2001), who found seasonal laminarinase
activity cycles in three Tasmanian mysid species, which imply a winter reliance on macroalgal
detritus. In summary, reliance on detritus of varied sources depends on species and perhaps on
location.
One point to emphasise is that most species of mysids, although they tend toward carnivory as
late juveniles and adults, do not appear to become exclusively carnivorous in the field (Hansson
et al. 1997, Branstrator 2000). Adults of some species, however, move very much toward that
extreme (Jerling & Wooldridge 1995, Kouassi et al. 2006). The vertically migrating species tend
to show increases in the fraction of detritus and sediments as gut contents during their benthic
phases (e.g., Lasenby & Shi 2004). On a diel timescale, they are sequential omnivores, alternating
between daytime diets rich in nitrogen and worth substantial digestive time and detrital diets that
yield higher net rate of gain at higher throughput rates (Zagursky & Feller 1985, Penry & Jumars
1987, Jumars 2000). Eating food of different qualities in sequence allows tailoring of gut retention
time to day-night change in diet quality (Penry & Jumars 1987, Jumars 2000), which can provide
much higher assimilation efficiency than optimising retention time on a complete mixture of daytime
with nighttime diets. Mysids in general have complex masticatory and digestive structures that not
only admit broad diets but also make ingestion of organisms that have no hard parts (e.g., many
protists and larval stages) difficult to identify (e.g., Brunet et al. 1994, De Jong-Moreau et al. 2001).
Mysids can be sufficiently abundant to influence prey community structure (e.g., Fulton 1983).
Both in the surf zone and in the plankton, larger diatom cells and chains experience the greatest
grazing pressure from mysids (e.g., Webb et al. 1987, 1988, Linden & Kuosa 2004). In a particular
interesting migratory role reversal from the typical offshore or estuarine scenario, Gastrosaccus
psammodytes Tattersall, 1958 feeds on surf-zone diatoms during the nighttime, benthic phase of
the diatoms (Webb et al. 1988), and Mysidopsis wooldridgei may consume up to 70% of surf-zone
diatom primary production (Webb & Wooldridge 1990). David et al. (2006b) found that juvenile
Mesopodopsis slabberi could account for the vast majority of herbivory in some settings. In fresh
water, daphnid populations can be severely reduced by mysids, defeating the purpose of mysid
introductions into lakes as food for salmonids (Spencer et al. 1999). In one shallow, tropical lagoon,
a population of mysids appeared to explain the low zooplankton:phytoplankton biomass ratio and
potentially could have consumed the entire holozooplankton production (Kouassi et al. 2006). In
that particularly carnivorous role, mysids can contribute substantially to phytoplankton nitrogen
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PETER A. JUMARS
requirements through excretion (Kouassi et al. 2006), but more typically the effect of feeding on
a combination of phytoplankton and zooplankton is expected to produce subtle shifts in size
structure of phytoplankton with little net effect on standing stocks (e.g., Linden & Kuosa 2004).
In estuaries, grazing pressure by mysids on holozooplankton varies among locations and seasons
but can be substantial (e.g., Fulton 1983, Aaser et al. 1995, Kibirige et al. 2003, Winkler et al.
2003). Early results with Americamysis bigelowi showed substantially reduced feeding on copepods
in the dark versus the light (Fulton 1982). Contrary to expectations from their well-developed eyes,
from the results of Fulton (1982) and from the observation by Clutter (1969) of more polarised
schooling in the light, more recent experimental data do not support the idea that most mysids are
primarily visual feeders. One migrating species showed higher ingestion rates on copepods in the
dark, whereas a non-migrating species showed no difference in light versus dark (Viherluoto &
Viitasalo 2001). Different mysid species apparently rely to differing degrees on tactile and visual
cues, and likely on chemical ones as well. It seems likely that species risking vertical migration
will have means to feed efficiently in the dark, that is, to obtain net gain during that migration risk
by ingesting large, lipid- and protein-rich prey. Moreover, mysids are capable of predation on
relatively large prey on the bottom as well (e.g., Wilhelm et al. 2002). One particularly interesting
observation is that, given a choice, some mysid species prefer meroplankton (David et al. 2006a).
Mysids thus have the potential to cause large mortality of meroplanktonic recruits in the plankton
or when they are settling. This potential coastal filter of larvae in both habitats (pelagic and benthic)
merits attention.
Habitat alteration and coupling
Roast et al. (2004) documented a significant role for mysids in resuspension of sediments underlying
the turbidity maximum of an estuary. Acoustic experiments with artificially emplaced sediments
(C.D. Jones & P.A. Jumars in preparation) in 20 m water depth off the Friday Harbor Laboratories
pier in the San Juan Islands, Washington, strongly implicated emergent Neomysis kadiakensis in
rapid microtopographic roughening of the sediment-water interface. Because the sediment surface
at steady state is already rough, this interaction of mysids is normally difficult to detect but likely
contributes to the erodibility effects documented by Roast et al. (2004), and attacks by predators
of mysids add further to it. Less obviously, mysids also have the potential to alter the lowermost
chemical boundary layer above the sediment-water interface, both through this roughening (e.g.,
Figure 1 of Jumars & Nowell 1984) as well as through movement over the bottom. The diffusive
sublayer in smooth-turbulent flow over the sea bed is typically on the order of 1 mm thick. Diffusion
time over that distance, calculated at time = distance2/(2D), where D is the molecular diffusion
coefficient (typically ≥ 2 × 10−9 m2 s−1), is 4 min. Thus disruption of the lowermost 1 mm every
4 min or less by mobile animals could appreciably increase benthic-pelagic fluxes by producing
unsteady diffusion through frequent sharpening of the diffusion gradient.
In terms of habitat coupling, diel migrations of holoplankton that feed closer to the surface
than where they spend other parts of the day result in net downward fluxes of nitrogen in the form
of excretion and carbon as respired dioxide (Longhurst & Harrison 1988, Longhurst et al. 1990)
in addition to the downward advection of particles as gut contents and faeces. Mysids returning
toward the seabed transport materials downward in all these ways. Unlike holoplankton, however,
emergent mysids also transfer material upward by feeding on the bottom and excreting and respiring
in surface waters. Net nitrogen flux is almost certainly downward because of the generally high
C:N ratios in benthic detrital foods, but where most carbon is obtained and carbon dioxide released
is less clear because swimming activity and temperature on average are both higher during the
pelagic phase. For materials highly concentrated in sediments, such as hydrophobic contaminants
and certain valence states and compounds of rare-earth elements and metals, mysid migrations and
112
food-web concentration conceivably could result in net upward transport (Evans et al. 1989, Neff
1997, Song & Breslin 1999).
Interesting potential exists for an interaction of habitat coupling and habitat modification by
dense, migratory swarms of mysids. Kunze et al. (2006) have documented a significant contribution
to turbulence and water-column mixing by coastal euphausiids during periods of stratification;
similar effects can be expected from mysids. One reason that diel migrants may be especially
effective at cross-isopycnal mixing is that, unlike shear-driven turbulence in the stratified ocean,
swimming is directed in the vertical, causing downward inertial jets during the upmigration and
vice versa.
A particularly tractable system for estimating, through daytime-collected egesta, the mysid
contribution to habitat coupling was identified by Carola et al. (1993), who worked with mysids
that migrate horizontally out of a submarine cave at night and return during the day, allowing
collection of faecal pellets in simple sediment traps set in the cave. Such submarine caves are shut
off from normal vertical particulate fluxes as well as being sheltered from normal horizontal
particulate fluxes through the obvious flow obstruction. They are generally considered oligotrophic.
Coma et al. (1997) estimated that the 756-m3 cave during the day housed 1–12 million mysids over
the seasons of their year-long study (peaking in spring). They estimated that the population’s daily
import to the cave through faecal deposition varied seasonally between 20 and 407 g dry wt of
particulate organic matter, 2–21 g C and 0.5–2.7 g N. Most pellets were released within 4.5 h after
the mysids’ sunrise return. The pellets are remarkably labile, with 20–50% of both C and N released
in dissolved form within 2 h after egestion, and they account for previous inabilities to reconcile
biological oxidation demand in such caves. Although it is a wonderful demonstration of intense
habitat coupling between the pelagic environment of the nearshore Mediterranean and a cave
environment, the results are difficult to generalise to sediment-dwelling mysids and their benthic-
pelagic coupling. The cave dwellers apparently do not feed during the day, and their nightly diet
remains obscure. Scanning electron microscopic examination of the pellets revealed no animal and
few phytoplankton remains, prompting Coma et al. (1997) to consider them to be detritivores, but
the lability of the pellets is then particularly puzzling. Possibilities compatible with the observed
detrital remains in the pellets are a diet rich in soft-bodied protists captured by feeding on marine
snow or rich in soft-bodied meroplankton (David et al. 2006b).
Dominating the holoplankton
Recent acoustic estimates of abundance call for renewed attention to mysid migrations. Two
geographically widely separated studies (Kringel et al. 2003, Taylor et al. 2005) found that the
water column during emergence was completely dominated by mysids, that is, that the mysid
contribution to nighttime standing stock exceeded daytime standing stocks of macrozooplankton
by an order of magnitude (Figure 3). Although widely separated, the environments share many
similarities. Both are in productive, shallow (10- to 20-m) regions of fjords that experience little
dilution by freshwater runoff, so whether and how deep these kinds of abundances extend over the
continental shelf is an obvious question. Another issue is whether acoustic methods would reveal
much higher abundances in those estuarine environments where high mysid abundances are already
known.
Midnight sinking was noted in many of the contour plots of abundance versus depth and time
made by Macquart-Moulin & Ribera Maycas (1995) for pelagic mysids. Midnight sinking in mysids
has been observed at least since the classic study of Neomysis americana in Narragansett Bay by
Herman (1963). Often but not always, mysids that emerge tend to be less abundant in near-surface
layers near the middle of the night than just after sunset or just before dawn. In some cases, a clear
mid-depth concentration accounts at least in part for this change (e.g., Herman 1963, the second
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PETER A. JUMARS
−30
A
Column scattering strength (dB)

−35
−40
−45
−50
0 12 0 12 0 12 0 12 0 12 0 12
Time of day (hours)
× 103
10
Sum of total biovolume (mm3 m−2)
B
8
3–6 m above bottom
0
0 12 0 12 0 12
Time of day (hours)
Figure 3 Changes in integrated water-column abundance derived from acoustic backscatter showing over-
whelming dominance by emergent animals. (A) Column scattering strength at 265 kHz from West Sound,
Orcas Island, Washington, in ~20 m of water. (From Kringel et al. 2003. With permission from the American
Society of Limnology and Oceanography, Inc.) Horizontal dotted lines connect median column scattering
strengths between and during emergence events, showing a 14-fold increase during emergence. The dashed
oval shows apparent midnight sinking (higher scattering strengths on both sides of a local minimum during
emergence. The first full day of the data record was 27 August 1995. (B) Column-integrated biovolumes
derived by acoustic inversion for the Damariscotta River estuary, mid-coast Maine, a shallow fjord. Bottom
depth is approximately 10 m, and the large emergence events that begin at the vertical dashed lines are cued
by the tides. (From Taylor et al. 2005, their Figure 3. With permission of the Estuarine Research Federation.)
Column-integrated standing stocks here also often display the molar- (tooth-) shaped structure with a local
minimum (dashed oval) and are 6-fold higher during emergence than during the day (Taylor et al. 2005). The
first full day of this record was 15 August 2002.
depth interval in the 4–5 April samples of his Figure 7; Macquart-Moulin & Ribera Maycas 1995,
bottom panel of their Figure 5 for Anchialina agilis). Anchialina agilis generally shows dusk and
dawn capture peaks in epibenthic sledge samples as well, but no evidence of a midnight return all
the way to the sea bed (Dauvin et al. 2000). Drivers of midnight sinking in mysids are not well known,
but satiety and predator evasion have been suggested in other taxa, and more recently midnight
sinking has been shown to promote retention of Calanus spp. in model results for one inland sea
114
(Emsley et al. 2005). It is worth remarking again that navigation through use of the polarised light
field should be most reliable both near the surface and near dawn and dusk, as one potential
contributor to surface congregation at those times.
Lending trophic and dynamic stability
Although they once were thought to have a destabilising effect on communities, recent and more
realistic, non-linear, non-equilibrium theory suggests an important stabilising role for omnivores
(McCann & Hastings 1997, Emmerson & Yearsley 2004) that has been confirmed in experiments
(Holyoak & Sachdev 1998). Omnivory is an obvious advantage when a favoured resource
becomes scarce, and multiple advantages accrue from eating one’s competitors for limiting
resources. Less obviously and more paradoxically, species that are eaten by many predators may
suffer lower total predation intensity (Montoya et al. 2006). The shelf ecosystems in which mysids
participate have high connectance (Link 2002, Dunne et al. 2004), but mysids have more connec-
tions over more trophic levels than does the average member of even those highly connected food
webs. In shallow water, mysids contribute in a major way to multichain omnivory, linking organic
matter that originates from benthic and pelagic primary production; theory originally developed
for lakes suggests that this linkage is particularly stabilising (Vadeboncoeur et al. 2005). By being
food for pelagic as well as benthic fishes and by consuming both planktonic and benthic primary
and secondary production and detritus, mysids not only link benthic and pelagic food webs but
also link resources with different inherent timescales. The latter linkages reduce synchrony and
overshoot in the populations constituting those resources, identifying another of the mechanisms
through which omnivores stabilise food webs (Rooney et al. 2006).
The northwest Atlantic shelf ecosystem has been disturbed at such high frequency and high
intensity for so long that it has not remained stable, with the notable crash of the groundfish fishery
(Link 2002, Choi et al. 2004). Although offshore banks have shown recovery of herring (Clupea
harengus) from about 1993 (Fogarty & Murawski 1998), some coastal spawning stocks have not
recovered from overfishing or environmental degradation (Smedbol & Stephenson 2001). Cascading
trophic effects from the groundfish collapse have been observed, with northern shrimp (Pandalus
borealis) and snow crab (Chionoecetes opilio) apparently experiencing release from predation
(Frank et al. 2005). A hint of the potentially stabilising effect of mysids comes from the conjecture
that they likely experienced a release from predation by benthic predators during the infamous cod
(Gadus morhua) crash in 1993 (Choi et al. 2004) and may have experienced some release from
herring predation as well. Juvenile cod captured in the southern Gulf of St. Lawrence shifted after
1990 from feeding on euphausiids to feeding on mysids, perhaps reflecting population expansion
in mysids, and as of 2000 had not shifted back (Hanson & Chouinard 2002).
Reasons why mysids have been underappreciated

Problems of sampling
The most obvious reason why mysids are not better understood is the difficulty of making total
areal (m−2) abundance estimates. It is a problem greatly compounded by simultaneous and (or)
sequential occupation of multiple habitats by mysids. Clutter & Anraku (1968) published a photo-
graph showing that information can be transmitted through a mysid school to turn it away from an
approaching plankton net. Lasenby & Sherman (1991) invoked the same explanation for greater
capture in vertically operated traps than in trawls. Fleminger & Clutter (1965) took oblique plankton
tows through a laboratory tank containing mysids (≥95% Metamysidopsis elongata) and captured
significantly more individuals in the dark than in the light. They noted, however, that they lacked
115
PETER A. JUMARS
observations to distinguish whether the difference was due to visual avoidance by the mysids or
to their potentially broader vertical distribution in the dark (the latter documented by Clutter 1969
in the field). Eleftheriou & Holme (1984) suggested that a 10% sampling efficiency is a reasonable
estimate for total macrofauna captured by epibenthic sledges. Emergence traps (e.g., Kringel et al.
2003) may also be biased to an unknown degree. Traps designed to catch emergent individuals
alter cues and may induce greater or lesser emergence than would occur in the absence of the trap.
Capture efficiency of emergent individuals is also unknown. Ribes et al. (1996) compared abundance
estimates based on faecal recovery in the aforementioned cave setting with net tows by divers and
found a 10-fold difference, with the net-based estimates being lower. A rare estimate of mysid
sampling efficiency based on independent sampling with several methods was that of Carlton &
Hamner (1987) from a coral reef lagoon. They found epibenthic sledge efficiency (relative to the
most efficient sampler, a trap operated by divers) for mysids to be between 1% and 10%. Because
nocturnally emerging mysids in coastal waters stay in the relative dark around the clock, often in
turbid waters, it is difficult to imagine how to undertake a complete census of a population analogous
to that carried out by Carlton & Hamner (1987).
Capture efficiency is certain to vary with life stage and species of mysid and likely with whether
they are schooling at the time of sampling, so the purpose of this short review of net and trap
sampling clearly is not to find or endorse a fixed estimate of sampling efficiency but rather to
indicate that most mysid sampling based on nets is likely to provide gross underestimates. One
underlying reason is high burst-swimming speed (caridoid escape reaction) and intermediate size
(order of 1 cm): fast nets in order to achieve reasonable filtration efficiencies typically use meshes
too large to capture mysids, whereas fine-meshed nets to achieve reasonable filtration efficiencies
are typically towed too slowly to capture mysids efficiently. For species that release eggs or weakly
swimming larvae into the plankton, slowly towed nets or plankton recorders will still reveal the
presence of large populations (e.g., Sameoto 2001). For mysids and other peracarids that brood
young, such gear will not sample adequately, especially if newly released juveniles initially are
benthic. When most places contain few individuals and a few places contain many mysids, a small
number of samples is unlikely to encounter the high concentrations of mysids. Although precise
estimates of mysid mean areal abundance would still be useful in evaluating their importance in
the ocean economy (e.g., in determining whether they must be included in estimates of secondary
production, nutrient regeneration and vertical fluxes), the more aggregated a population is, the less
relevance mean abundances have to issues such as encounter rates and behaviours (Omori & Hamner
1982). Direct observations by divers find mysid swarms up to densities of nearly 600 l−1 (6 × 105 m–3).
Such dense swarms are usually dominated by a single species but may be joined by several (Ohtsuka
et al. 1995). Instead of focusing on mean abundance it may make more sense to be concerned about
the fraction of space occupied by swarms and the frequency with which any given place is occupied
by a swarm.
A simple exercise demonstrates several aspects of the mysid-sampling problem. For Anchialina
agilis, because the entire population appears to emerge at night, it would appear to be feasible to
estimate total areal abundance by vertically integrating depth-specific abundance estimates. For the
points near 2020 h on 11 December for male A. agilis in the 60-m water column as depicted by
Macquart-Moulin & Ribera Maycas (1995, their Figure 4) integrating by linear interpolation
between the four depths sampled yields an areal abundance of ~15.4 males m−2. For females plus
juveniles on the same night, two sections are plotted (their Figure 5) that show peak abundances
at about 0500 h of ~3 and ~15.4 ind. m−2, respectively, for an average areal estimate for females and
juveniles of ~9 ind. m−2. The sum (total of males plus females and juveniles) exceeds typical,
sledge-estimated abundances for the species, and itself is likely an underestimate because of net
avoidance by pelagic A. agilis, but could be an overestimate of local benthic areal abundance during
the day if there is significant local horizontal convergence during emergence.
116
Acoustic estimates of abundance based on vertically integrated biovolumes and of migration rates
based on changes in vertically resolved biovolumes are also underestimates of unknown severities.
A continuing issue with acoustic estimates of organism abundance is individual target strength,
particularly for elongate organisms with complex morphologies, because their target strengths vary
with orientation relative to the transducer, greatly increasing the uncertainty of abundance estimates
made through acoustic inversion (e.g., Amakasu & Furusawa 2005, Lawson et al. 2006). Time-
varying polarizations and direction changes of swarms and schools clearly compound this problem,
but acoustic methods are more robust in biological application than some of the complexities in
acoustic inversion might suggest (Benoit-Bird & Au 2002).
A more fundamental problem, often opined (Pearre 2003), is that methods that infer migration
from changes in spatial distributions are unable to detect migrations that do not cause net change
(Figure 4). Thus it is not known to what extent acoustic observations of Neomysis kadiakensis (e.g.,
Kringel et al. 2003) or N. americana (e.g., Abello et al. 2005, Taylor et al. 2005) reflect individuals
staying in the water column for hours at a time versus frequently exchanging places with benthic
members of the same population, and it is not known what fraction of the total population fails to
migrate at all. Mysid species and life stages differ in this regard, as reflected in near-surface and
near-bottom or nearshore and offshore samples taken at different times of day. Acoustic methods
based on backscatter intensity in principle could be used to obtain data fairly close to the bottom
by burying an upward-looking device. Besides the logistical problems, however, most transducers
have non-ideal near-field effects, and TAPS (Tracor Acoustic Profiling System, now under the
auspices of BAE Systems, San Diego, California) on this account does not collect data closer than
37.5 cm from its transducers. Conversely, down-looking geometries (e.g., Greenlaw et al. 2004)
are effectively ‘blinded’ in the lowermost one or two range bins by the high backscatter intensity
from the bottom. Use of split-beam, dual-beam and multibeam acoustics, generating the acoustic
equivalent of binocular vision and giving some capability of tracking individuals, promises at long
last to reveal individual zooplankton behaviours (Smith et al. 1992, Kaufmann et al. 1993,
De Robertis et al. 2003). These methods do not overcome interference from strong bottom scatter-
ing, but they should allow individual acoustic target strengths and trajectories of individuals to be
determined from stable bottom platforms, at least when current speeds are small and acoustic
sampling intervals are short. Each sampling method will need to be used for its strengths and to
remedy the weaknesses of others. Acoustics have the clear advantage in spatial and temporal
resolution (e.g., Figure 5), but have the generic problem of being poor at resolving identities. When
species diversity is low, however, acoustic methods can be sufficient for discrimination (David et al.
1999), and shape and biomechanical properties that are detectable acoustically show promise of
providing better discrimination in the future. Acoustic methods excel with their high resolution in
both space and time at pinpointing locations and times of interest for sampling by other means
(e.g., Figure 6).
Difficulty in modelling
Whereas high, flow-driven horizontal flux over the bottoms they inhabit is an important component
of the ‘umwelt’ of mysids that emerge in coastal environments, it compounds the difficulties of
simulation. It is no accident that many of the citations given here are for studies done in closed
systems (lakes) and semi-enclosed systems (estuaries) where effects are much easier to calculate
and model, rather than the nominal coastal environments of primary interest. Open-shelf systems
are more difficult to constrain in both measurements and models. Further, the same infidelity to
habitat that makes abundance estimates difficult also complicates study in analog laboratory systems.
Although smaller tanks have yielded valuable observational data on taxa without extensive migra-
tions (Fosså 1986), a much larger system (e.g., Conover & Paranjape 1977) would be needed to
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PETER A. JUMARS
A B
3/4 5/8 5/8 5/8 5/8 5/8 5/8 5/8 3/4 6 5 4 3 2 3 4 5 6
5/8 1/2 1/2 1/2 1/2 1/2 5/8 5/8 3/4 5 4 3 2 1 2 3 4 5
5/8 1/2 1/2 5/8 5/8 3/4 After midnight p toward 4 3 2 1 0 1

source = 4 × p away from source
3/4 5/8 3/4 Before midnight p = 1/8 for all steps 5 4 3 Smallest number of steps from source
T=1h T = 12 h
(6 time steps) (72 time steps)
T=2h T = 11 h
T=3h T = 10 h
C
T=4h T=9h
T=5h T=8h
T=6h T=7h
0.05 0.1 0.15 0.2 0.25 0.3 0.35 0.4

Fraction of individuals in that compartment
Figure 4 A simple demonstration of the potential for radically different mysid behaviours to produce similar patterns of
vertical change in abundance over large regions of the habitat and the consequent need to develop methods for tracking
individuals during migration. Simple, heuristic Markov models (cf. Kemeny & Snell 1960, Jumars et al. 1981) were set
up under two different sets of hypothetical behaviours. The boxes in panels A and B represent a 2-dimensional section
in the onshore–offshore direction, shoaling to the right. Greyed boxes (A, B) represent a hyperbenthic environment, with
white boxes being in the water column. In both model runs (see also next page), all animals began in the box labelled 0
in panel B. Time steps were 10 min. In the first run, all transition probabilities were set to 1/8; that fraction of individuals
in each box moved to the adjacent box, and the fractions in (A) represent the proportion that stayed in the box during
one time step. After a long time with this transition matrix, individuals would become uniformly distributed. At T = 6 h,
however, the original source box became an absorbing state (transitions out were prohibited) representing recognition of
the habitat by the model animals. In addition, all transition probabilities that reduced the number of steps back to the
original source box were made five times larger than those leading away. (Continued on next page.)
118
T=1h T = 12 h
T=2h T = 11 h
T=3h T = 10 h
D
T=4h T=9h
T=5h T=8h
T=6h T=7h
0.05 0.1 0.15 0.2 0.25 0.3 0.35 0.4

Fraction of individuals in that compartment
Figure 4 (continued) That process led to the return of 95% of all individuals in 6 h. This first scenario can be construed
as a broadening of the depth distribution through very active random transfer among compartments, with a biased
random walk bringing the animals back. Arrows in panel C represent the cycle that emergence would follow from start
to finish. If the cycle is repeated (dashed arrow), results of the slightly different starting condition are not perceptibly
different. In the second scenario (D) the same fractions remained in each compartment as shown in (A). Initially, all
transitions moving a greater number of steps away from the same source box were set at 1/20 (with zero probability
of moving toward the source), except that downward transitions were ruled out (also set at zero) and the model was
run for 6 h before reversal of behaviour (moving away from the original source set to zero probability and moving
toward set to 4/25). In this phase, the initial source compartment was again made perfectly absorbing. The second
scenario can be construed as much slower, deliberate navigation to and from nighttime feeding areas. Because net
movement among boxes is not radically different between the two scenarios (except for deep regions in panel D that
are effectively unreachable), neither is the depth-time distribution of individuals. Again 95% of individuals returned
after 72 time steps, and running the cycle again with this starting condition produces results that are not perceptibly
different from starting all individuals in the original source compartment. Animals in the first scenario do a great deal
more moving among locations per unit of time and therefore would likely have much higher encounter rates with
predators. Future methods need to differentiate net from gross migration rates. Note that while neither scenario includes
any sinking at all, the pattern of abundance in the surface compartment above the source shows what might be interpreted
as midnight sinking, that is, higher abundances early and late in the migration cycle than in its middle.
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PETER A. JUMARS
Day of July 1995, West Sound, Orcas Island, Washington

26 27 28 29
Height above TAPS (m)
20
10
0
midnight noon midnight noon midnight noon
−75 −70 −65 −60 −55 −50 −45 −40 −35 −30 −25
Volume backscattering strength at 265 kHz (dB)
Day of August 2002, Damariscotta River, Maine

19 20 21 22
10
0
midnight noon midnight noon midnight
0 200 400 600 800 1000 1200 1400 1600

Biovolume concentration (mm3 m−3)
Figure 5 (See also Colour Figure 5 in the insert following page 344.) Resolution achievable acoustically
without resolving individual mysids, showing some of the diversity that can be encountered in emergence. The
two time series have been aligned so that midnight occurs in both at the same location on the abscissa. Regularity
of nightly emergence in the West Sound time series (formed from data taken every 2 min in 25-cm range bins;
top panel from Kringel et al. 2003. With permission of the American Society of Limnology and Oceanography,
Inc.) is stunning, and the descents are often more organised than the initial ascent, a phenomenon herein called
the Dracula effect (Abello et al. 2005). The data from the Damariscotta estuary have even higher resolution
(taken every minute with 12.5-cm spatial resolution). Low tides occurred near noon in the West Sound series,
and the reflection from the water surface is evident as bright red patches of high backscatter. The Damariscotta
time series shows both low tides of the day, with the red extensions from the surface being a combination of
fish aggregations and bubbles injected by breaking wavelets from the afternoon sea breeze. The Damariscotta
series shows multiple emergence events per night. The first emergence events after dark are light cued (Abello
et al. 2005), whereas the largest events begin at peak tidal flow speed after dark (Taylor et al. 2005). (Bottom
panel from Taylor et al. 2005. With permission of the Estuarine Research Federation.)
Figure 6 (opposite page; see also Colour Figure 6 in the insert.) Acoustic methods that do not resolve
individual mysids can also be used to gather information on group ascent and descent velocities and times of
departure from and return to the sea bed. (After Kringel et al. 2003, from the same 1995 dataset presented at
the top of Figure 5. With permission of the American Society of Limnology and Oceanography.) Leading
edges of the ascent can be recognised as the first and the last pixels above background backscattering levels
at each height above bottom (white circles). Slopes of the linear regressions yield ascent and descent velocities,
and intercepts provide objective start and stop times of emergence. Modal abundance values (blue X marks,
colour version) represent the behaviours of a larger fraction of the emergent population, but again only net
migration rates can be visualised from measures of total backscatter.
120
20 20 20
15 15 15
−25
10 10 10
5 5 5
−30
20 22 24 2 20 22 24 2 20 22 24 2
26–27 July 27–28 July 28–29 July
−35
20 20 20
15 15 15 −40
10 10 10
Volume backscattering strength (dB)

5 5 5 −45
20 22 24 2 20 22 24 2 20 22 24 2
29–30 July 30–31 July 30 July–1 August −50
20 20 20
15 15 15 −55
10 10 10
−60
5 5 5
2 4 6 8 2 4 6 8 2 4 6 8
27 July 28 July 29 July −65
20 20 20
−70
15 15 15
10 10 10
−75
5 5 5
2 4 6 8 2 4 6 8 2 4 6 8
30 July 31 July 1 August
Hour of day
Figure 6 (See facing page for caption.)
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PETER A. JUMARS
provide realistic prey distributions and allow migration. Moreover, mysids are notoriously sensitive
to light exposure and take considerable time to recover physiologically, and presumably behav-
iourally, from unnatural levels (e.g., Attramadai et al. 1985, Lindström et al. 1988). Red-light
viewing to avoid light-induced artifacts (e.g., Fosså 1986) rapidly becomes less practical at long
range because red light has a high absorption coefficient in water.
Challenges and opportunities

A continuing challenge is to produce an effective, convincing census of shelf mysid populations
(abundance versus location versus time) that misses neither migrating nor non-migrating members.
No one method yet attempted appears to be a panacea in this regard because different methods
have different strengths and weaknesses in different habitats.
In this review, emphasis has been placed on the challenges in measuring the vertical component
of migration and the coupling of benthic with pelagic habitats because enough data can be gathered
to review them. The horizontal component provides many additional challenges and opportunities.
It is obviously a key feature of mysid migrations in and out of caves (e.g., Carola et al. 1993, David
et al. 1999) and vegetation (e.g., Wittman 1977, Mattila et al. 1999). Whereas organisms with clear
structural, spatial refuges from predation can return to them when risk is high, organisms without
a structural spatial refuge adopt other means of reducing risk. Besides the obvious tactic of staying
in the dark, emergent mysids can alter schooling patterns and (or) reduce their feeding rates to
reduce risk by evading pursuit or detection, respectively (Ritz 1994, Hamren & Hansson 1999).
Mysids are documented to change behaviour via both visual and chemical cues from predators
(Hamren & Hansson 1999, Cohen & Ritz 2003, Linden et al. 2003). Schooling for mysids that
have evolved schooling behaviours may be more effective at predator evasion than is their entry
into vegetation because such entry breaks up schools into more easily taken, smaller groups (Flynn
& Ritz 1999), and swarm-formers versus species using structural refugia show the expected
responses (Linden et al. 2003). Burrowing or burying during the day by emergent mysids appears
to be an intermediate solution, providing mechanical, optical and perhaps chemical refuge from
sensory detection by predators, but it probably has costs of reduced feeding capabilities relative to
mysids feeding at the sediment-water interface. Mysid schooling may be less effective against
predators that themselves school (Foster et al. 2001) and may incur greater risk from suspension-
feeding animals that do not pursue individual prey but filter them in bulk, such as gray whales
(Eschrichtius robustus). School size appears to depend on a dynamic optimum solution between
reduction of risk and enhancement of feeding success (Ritz 1994). Among questions that merit
attention are whether mysids by schooling during emergence and re-entry enhance their mean
individual rates of capture of emergent and holoplanktonic copepods and whether those in poorest
nutritional status take greater risks by emerging earlier in the night as has been demonstrated for
some diel migrators among the holoplankton (Hays et al. 2001).
Near shore, where physical gradients in nearly all directions and over time are usually strong,
gradients in risk and potential gain run in all directions but certainly covary strongly with distance
from shore, water depth and time of day (e.g., Takahashi et al. 2004). Diel horizontal migration
has not received much recent attention in shallow-water biological oceanography, whereas diel
horizontal differences in abundance are so commonly documented among freshwater crustaceans
and rotifers that the term ‘shore avoidance’ was coined when the behaviour of swimming toward
open water in the morning and a putative orientation mechanism in the form of polarised light were
reported (Siebeck 1968a,b). Curiously, marine mysid capability to detect polarisation was docu-
mented a decade earlier (Bainbridge & Waterman 1957), but early marine work focused more on
physiology than on ecological implications. More recently, Goddard & Forward (1991) documented
122
grass shrimp orientation mediated by polarisation and aiming them away from shore during daylight,
likely avoiding littoral predators. Test results are consistent with several predictions based on
predator avoidance in fresh waters (e.g., Wicklum 1999). Recent experimental verification of
homing capabilities in marine mysids (Twining et al. 2000) and clarification of submarine polari-
sation showing potential utility for orientation to >200 m depth (Waterman 2005) promise renewed
interest. Notably, horizontal tilt of the polarisation e-vector is greatest near dawn and dusk, giving
the best onshore–offshore information then. Benefits of homing to particular reef, cave or macro-
phyte sites are obvious, but information on relocation of mysids from horizontally (alongshore)
more uniform environments is scarce. Wittman (1977), however, painted mysids in his diving study
and remarked that he could find them up to 12 days later (the intermolt period for one of the
dominant species). Further mark-recapture results might be particularly informative about homing
and might also be revealing about population sizes if the relevant parameters can be estimated.
In terms of risks, many also vary over timescales beyond the diel. Over 6 yr of observation,
we have observed Neomysis americana to be present in our local, shallow fjord (Damariscotta
River estuary, with little freshwater input) weeks before any appreciable vertical migration begins.
Nocturnal emergence does not start until late May or early June, coincident with the termination
of the annual alewife (Alosa pseudoharengus) run upstream (P.A. Jumars, personal observations).
A mechanistic relationship remains to be established, but a time-varying predation risk at mid-
latitudes from seasonal cycles of clupeids is nearly universal.
In terms of spatiotemporal gradients of predators, recent acoustic results document that, in a
setting with a narrow shelf, many mid-water animals migrate strikingly close to shore at night,
with onshore–offshore speeds exceeding those of their vertical excursions by three orders of
magnitude (Benoit-Bird et al. 2001, Benoit-Bird & Au 2004). Although wider shelves will impede
mid-water participants in diel migration, analogous phenomena in diel predator movemements are
expected even on broad shelves. For emergent mysids that time-share food resources and predation
risks between pelagic and benthic habitats, the likelihood that optimal benthic and pelagic habitats
occur in the same depth of water seems remote. What is perhaps the greatest animal migration on
the planet has recently been revealed acoustically and comprises horizontal movements of great
shoals of fishes over the continental shelf (Makris et al. 2006); diel patterns are not yet established.
It is easy to predict with confidence that the horizontal and coastal equivalent of Vinogradov’s
ladder (Vinogradov 1962) will be documented in the observatory era of oceanography and that
mysids will constitute several rungs.
Combined gradients in predation risk and resources can produce confusing patterns of resource
use when interpreted according to resource availability alone, particularly for omnivores that have
multiple feeding options (Morris 2005). In an innovative and ambitious contribution, Speirs et al.
(2002) predicted and dissected the movements of intertidal mysids as functions of both resource
and predator effects and time, through population-level modelling of distributions. It could be useful
to repeat their measurements of response to sediment organic quality using a method more sensitive
to the labile, digestible component of sediments (Mayer et al. 1995), which in general is not well
correlated with total organic carbon (Dauwe et al. 1999) and using only the surficial and resus-
pendable materials (Mayer et al. 1993) to which epibenthic mysids are exposed.
A quickly advancing, alternative approach to modelling organisms that use multiple habitats
also appears to merit closer examination for potential application to mysids. Given the long history
of research on diel vertical migration in zooplankton, prospects for rapid advance to predictive
understanding of emergence by reductionist testing of the same or variant collections of hypotheses
on mysids or other emergers appear bleak. As an alternative, IBM is a subset of agent-based
modelling within complexity theory (Auyang 1998) and has several characteristics that distinguish
it from classical reductionist or holistic approaches (Grimm & Railsback 2005, p. 55):
123
PETER A. JUMARS
• “Theory is neither holist (system-level) nor reductionist (individual-level). We do not

assume that ecological systems can be understood from only the system level, but we
also do not assume that a system is simply the sum of its individual parts. Systems have
properties of completely different types than the properties of individuals, and theory
must explain these system properties.
• Theory must therefore be multilevel, linking traits of individuals to properties of the
system. We are not interested in understanding all aspects of individual behavior but
instead are interested in developing models of individuals that explain important system
properties.
• Observational and experimental science at both the individual and system level is the
basis for theory development. Such empirical science is important both for discovering
the phenomena driving the system and for testing theories.”
IBM is also an obvious approach toward understanding of emergence because the constellation
of traits associated with emergent mysids overlaps so broadly with published success stories of
IBM in explaining and predicting schooling and foraging behaviours under varying risks, dispersal,
habitat usage and local reproductive success (Grimm & Railsback 2005, Chapter 6). What is
particularly promising about this approach is that it frequently predicts very different consequences
in different environments, as would appear necessary in the case of Neomysis americana. IBMs
have already been used in other marine applications (Miller et al. 1998, Grimm et al. 1999, Crain
& Miller 2001, Leising 2001). The variety of IBM that would appear appropriate to mysids assumes
that individuals choose behaviours that on average enhance their fitness, and those behaviours are
termed ‘adaptive traits’ (Zhivotovsky et al. 1996). A successful IBM is generally recognised through
correct prediction of often-subtle spatial patterns of distribution and habitat usage (Dieckmann et al.
2000, Grimm & Railsback 2005). Both from the standpoint of understanding observations and
making models, the words of Pearre (2003) resonate: “Without knowing the actual movements
of individuals it seems unlikely that we will be able to understand their causes, nor the effects of
vertical migrations on the environment or on the migrators themselves”.
Acknowledgements
I am grateful to the Office of Naval Research (grant N00014-03-1-0776) for generous funding of
my acoustic research on mysid migrations and to Van Holliday and Charles Greenlaw of BAE
Systems for perpetual help and encouragement with acoustic approaches. Much of this synthesis
comes from the work of four master of science students: Kelly Kringel, Heather Abello, Leslie
Taylor and Mei Sato. The manuscript benefitted considerably from comments by K. Dorgan and
careful editing by Prof. R.J.A. Atkinson. I thank Dennis Allen and Harmon Brown for sharing their
data and ideas about Neomysis americana in South Carolina and on Georges Bank, respectively.
References
Aaser, H., Jeppesen, E. & Sondergaard, M. 1995. Seasonal dynamics of the mysid Neomysis integer and its
predation on the copepod Eurytemora affinis in a shallow hypertrophic brackish lake. Marine Ecology
Abello, H.U., Shellito, S.M., Taylor, L.H. & Jumars, P.A. 2005. Light-cued emergence and re-entry events in
a strongly tidal estuary. Estuaries 28, 487–499.
Albertsson, J. 2004. Trophic interactions involving mysid shrimps (Mysidacea) in the near-bottom habitat in
the Baltic Sea. Aquatic Ecology 38, 457–469.
124
Allen, D.M. 1984. Population dynamics of the mysid shrimp Mysidopsis bigelowi W.M. Tattersall in a
temperate estuary. Journal of Crustacean Biology 4, 25–34.
Amakasu, K. & Furusawa, M. 2005. The target strength of Antarctic krill (Euphausia superba) measured by
the split-beam method in a small tank at 70 kHz. ICES Journal of Marine Science 63, 36–45.
Antezana, T. 1999. Plankton of southern Chilean fjords: trends and linkages. Scientia Marina 63 (Suppl. 1),
69–80.
Apel, M. 1992. Spatial distribution and seasonal occurrence of Mysidacea in the Jade estuary (North Sea,
Germany), with some comments on diurnal migrations. In Taxonomy, Biology and Ecology of (Baltic)
mysids (Mysidacea: Crustacea), J. Köhn et al. (eds). Rostock, Germany: Rostock University Press,
98–108.
Ariani, A.P., Wittmann, K.J. & Franco, E. 1993. A comparative study of static bodies in mysid crustaceans:
evolutionary implications of crystallographic characteristics. Biological Bulletin (Woods Hole) 185,
393–404.
Aronson, R.B. 1989. The ecology of Octopus briareus Robson in a Bahamian saltwater lake. American
Malacological Bulletin 7, 47–56.
Astthorsson, O.S. 1985. Mysids occurring in the stomachs of cod caught in the Atlantic water south and west
of Iceland. Sarsia 70, 173–178.
Attramadai, Y.G., Fosså, J.H. & Nilsson, H.L. 1985. Changes in behaviour and eye-morphology of Boreomysis
megalops G.O. Sars (Crustacea: Mysidacea) following exposure to short periods of artificial and
natural daylight. Journal of Experimental Marine Biology and Ecology 85, 135–148.
Audzijonyte, A. & Väinölä, R. 2005. Diversity and distributions of circumpolar fresh- and brackish-water
Mysis (Crustacea: Mysida): descriptions of M. relicta Lovén, 1862, M. salemaai n. sp., M. segerstralei
n. sp. and M. diluviana n. sp., based on molecular and morphological characters. Hydrobiologia 544,
89–141.
Auyang, S.Y. 1998. Foundations of Complex System Theories in Economics, Evolutionary Biology and
Statistical Physics. New York: Cambridge University Press.
Azeiteiro, U.M.M., Jesus, L. & Marques, J.C. 1999. Distribution, population dynamics, and production of the
suprabenthic mysid Mesopodopsis slabberi in the Mondego Estuary, Portugal. Journal of Crustacean
Biology 19, 498–509.
Bacescu, M. 1941. Les Mysidacés des eaux Méditerranéennes de la France (spécialment de Banyuls) et des
eaux de Monaco. Bulletin de l’Insitut Océanographique (Monaco) 795, 1–46.
Bacescu, M. & Schiecke, U. 1974. Gastrosaccus magnilobatus n. sp., and Erythrops peterdohrni n. sp.
(Mysidacea) — new surprises from the Mediterranean. Crustaceana 27, 113–118.
Bainbridge, R. & Waterman, T.H. 1957. Polarized light and the orientation of two marine Crustacea. Journal
of Experimental Biology 34, 342–364.
Bainbridge, R. & Waterman, T.H. 1958. Turbidity and polarized light orientation of the crustacean Mysidium.
Journal of Experimental Biology 35, 487–493.
Bamber, R.N. & Henderson, P.A. 1994. Seasonality of caridean decapod and mysid distribution and movements
within the Severn Estuary and Bristol Channel. Biological Journal of the Linnean Society 51, 83–91.
Barber, R.T. & Smith, W.O., Jr. 1981. The role of circulation, sinking, and vertical motion in sorting of
phytoplankton in the upwelling center at 15S. In Coastal Upwelling, F.A. Richards (ed.). Washington,
D.C.: American Geophysical Union, 366–371.
Beaudouin, J. 1979. Euphausiaces, mysidaces, larves de decapodes du Golfe de Gascogne (plateau continental).
Revue des Travaux de l’Institut des Pêches Maritimes 43, 367–389.
Benoit-Bird, K.J. & Au, W.W.L. 2002. Energy: converting from acoustic to biological resource units. Journal
of the Acoustical Society of America 111, 2070–2075.
Benoit-Bird, K.J. & Au, W.W.L. 2004. Diel migration dynamics of an island-associated sound-scattering layer.
Deep-Sea Research 51, 707–719.
Benoit-Bird, K.J., Au, W.W.L., Brainard, R.E. & Lammers, M.O. 2001. Diel horizontal migration of the
Hawaiian mesopelagic boundary community observed acoustically. Marine Ecology Progress Series
217, 1–14.
Beyst, B., Buysse, D., Dewicke, A. & Mees, J. 2001. Surf zone hyperbenthos of Belgian sandy beaches:
seasonal patterns. Estuarine, Coastal and Shelf Science 53, 877–895.
125
PETER A. JUMARS
Boudreau, B.P. & Jørgensen, B.B. (eds) 2001. The Benthic Boundary Layer: Transport Processes and Bio-
geochemistry. Oxford: Oxford University Press.
Bozzano, A., Recasens, L. & Sartor, P. 1997. Diet of the European hake Merluccius merluccius (Pisces:
Merluciidae) in the western Mediterranean (Gulf of Lions). Scientia Marina 61, 1–8.
Brandt, A., Linse, K. & Weber, U. 1997. Abundance and diversity of peracarid taxa (Crustacea, Malacostraca)
along a transect through the Beagle Channel, Patagonia. Polar Biology 18, 83–90.
Branstrator, D.K. 2000. Measuring life-history omnivory in the opossum shrimp, Mysis relicta, with stable
nitrogen isotopes. Limnology and Oceanography 45, 463–467.
Brattegard, T. & Meland, K. 1997. Mysidacea (Crustacea) in the Faroe area. Fródskaparrit 45, 69–95.
Brierley A.S., Brandon M.A. & Watkins J.L. 1998. An assessment of the utility of an acoustic Doppler current
profiler for biomass estimation. Deep-Sea Research I 45, 1555–1573.
Brown, H., Bollens, S.M., Madin, L.P. & Horgan, E.F. 2005. Effects of warm water intrusions on populations
of macrozooplankton on Georges Bank, northwest Atlantic. Continental Shelf Research 25, 143–156.
Brunel, P. 1979. Seasonal change of daily vertical migrations in a suprabenthic cold-layer community over
mud in the Gulf of St. Lawrence. In Cyclic Phenomena in Marine Plants and Animals, E. Naylor &
R.G. Harnoll (eds). Oxford: Pergamon Press, 383–390.
Brunet, M., Arnaud, J. & Mazza, J. 1994. Gut structure and digestive cellular processes in marine Crustacea.
Buchanan, J.B. & Hedley, R.H. 1960. A contribution to the biology of Astrorhiza limicola (Foraminifera).
Journal of the Marine Biological Association of the United Kingdom 39, 549–560.
Buhl-Jensen, L. & Fosså, J.H. 1991. Hyperbenthic crustacean fauna of the Gullmarfjord area (western Sweden):
species richness, seasonal variation and long-term changes. Marine Biology 109, 245–258.
Calliari, D., Cervetto, G., Gomez, M. & Bastreri, D. 2001. Neomysis americana from Solis Grande, Uruguay.
Atlântica, Rio Grande 23, 117–125.
Carlton, J.H. & Hamner, W.M. 1987. A diver-operated device for capture of mobile epibenthic organisms.
Limnology and Oceanography 32, 503–510.
Carola, M., Coma, R., Riera, T. & Zabala, M. 1993. Fecal pellets collection as a method for assessing egesta
of the marine cave-dwelling mysid Hemimysis speluncola. Scientia Marina 57, 51–63.
Carter, J.H.C. & Dadswell, M.J. 1983. Seasonal and spatial distribution of planktonic Crustacea in the lower
Saint John River, a multibasin estuary in New Brunswick, Canada. Chesapeake Science 6, 142–153.
Cartes, J.E. & Sorbe, J.C. 1995. Deep-water mysids of the Catalan Sea: species composition, bathymetric and
near-bottom distribution. Journal of the Marine Biological Association of the United Kingdom 75,
187–197.
Castel, J. 1993. Long-term distribution of zooplankton in the Gironde estuary and its relation with river flow
and suspended matter. Cahiers de Biologie Marine 34, 145–163.
Champalbert, G. & Macquart-Moulin, C. 1970. Les Péracarides de l’hyponeuston nocturne du Golfe de
Marseille. Cahiers de Biologie Marine 11, 1–29.
Chess, D.W. & Stanford, J.A. 1999. Experimental effects of temperature and prey assemblage on growth and
lipid accumulation by Mysis relicta Loven. Hydrobiologia 412, 155–164.
Choi, J.S., Frank, K.T., Leggett, W.C. & Drinkwater, K. 2004. Transition to an alternate state in a continental
shelf ecosystem. Canadian Journal of Fisheries and Aquatic Sciences 61, 505–510.
Clutter, R.I. 1967. Zonation of nearshore mysids. Ecology 48, 200–208.
Clutter, R.I. 1969. The microdistribution and social behaviour of some pelagic mysid shrimps. Journal of
Clutter, R.I. & Anraku, M. 1968. Avoidance of samplers. In Zooplankton Sampling, D.J. Tranter (ed.). Paris:
UNESCO, 57–76.
Cohen, P.J. & Ritz, D.A. 2003. Role of kairomones in feeding interactions between seahorses and mysids.
Colman, J.S. & Segrove, F. 1955. The tidal plankton over Stoupe Bank Sands, Robin Hood’s Bay. Journal of
Animal Ecology 24, 445–462.
Coma, R., Carola, M., Riera, T. & Zabala, M. 1997. Horizontal transfer of matter by a cave-dwelling mysid.
Pubblicazioni della Stazione Zoologica di Napoli I: Marine Ecology 18, 211–226.
126
Conover, R.J. & Paranjape, M.A. 1977. Comments on the use of a deep tank in planktological research.
Helgoland Marine Research 30, 105–117.
Cornet, M., Lissalde, J.-P., Bouchet, J.-M., Sorbe, J.C. & Amoureux, L. 1983. Données qualitatives sur le
benthos et le suprabenthos d’un transect du plateau contiental sud-Gascogne. Cahiers de Biologie
Marine 24, 69–84.
Cowles, R.P. 1930. A biological study of the offshore waters of Chesapeake Bay. Bulletin of the United States
Bureau Fisheries 46, 277–381.
Crain, J.A. & Miller, C.B. 2001. Effects of starvation on intermolt development in Calanus finmarchicus
copepodites: a comparison between theoretical models and field studies. Deep-Sea Research II 48,
551–566.
Cunha, M.R., Sorbe, J.C. & Bernardes, C. 1997. On the structure of the neritic suprabenthic communities
from the Portuguese continental margin. Marine Ecology Progress Series 157, 119–137.
Cunha, M.R., Sorbe, J.C. & Moreira, M.H. 1999. Spatial and seasonal changes of brackish peracaridan
assemblages and their relation to some environmental variables in two tidal channels of the Ria de
Aveiro (NW Portugal). Marine Ecology Progress Series 190, 69–87.
Darling, J.D., Keogh, K.E. & Steeves, T.E. 1998. Gray whale (Eschrichtius robustus) habitat utilization and
prey species off Vancouver Island, British Columbia. Marine Mammal Science 14, 692–720.
Dauby, P.A. 1995. A ∂13C study of the feeding habits in four Mediterranean Leptomysis species (Crustacea:
Mysidacea). Pubblicazioni della Stazione Zoologica di Napoli I: Marine Ecology 16, 93–102.
Dauvin, J.-C. & Vallet, C. 2006. The near-bottom layer as an ecological boundary in marine ecosystems:
diversity, taxonomic composition and community definitions. Hydrobiologia 555, 49–58.
Dauvin, J.-C., Vallet, C., Mouny, P. & Zouhiri, S. 2000. Main characteristics of the boundary layer macrofauna
in the English Channel. Hydrobiologia 426, 139–156.
Dauwe, B., Middelburg, J.J., Van Rijswijk, P., Sinke, J., Herman, P.M.J. & Heip, C.H.R. 1999. Enzymatically
hydrolyzable amino acids in the North Sea sediments and their possible implication for sediment
nutritional values. Journal of Marine Research 57, 109–134.
David, P.M., Guerin-Ancey, O. & Van Cuyck, J.P. 1999. Acoustic discrimination of two zooplankton species
(mysid) at 38 and 120 kHz. Deep-Sea Research 46, 319–333.
David, V., Chardy, P. & Sautour, B. 2006a. Fitting a predator-prey model to zooplankton time-series data in
the Gironde estuary (France): ecological significance of the parameters. Estuarine, Coastal and Shelf
Science 67, 605–617.
David, V., Sautour, B., Chardy, P. & Leconte, M. 2005. Long-term changes of the zooplankton variability in
a turbid environment: the Gironde estuary (France). Estuarine, Coastal and Shelf Science 64, 171–184.
David, V., Sautour, B., Galois, R. & Chardy, P. 2006b. The paradox [of] high zooplankton biomass — low
vegetal particulate organic matter in high turbidity zones: what way for energy transfer? Journal of
Dean, A.F., Bollens, S.M., Simenstad, C. & Cordell, J. 2005. Marshes as sources or sinks of an estuarine
mysid: demographic patterns and tidal flux of Neomysis kadiakensis at China Camp marsh, San
Francisco estuary. Estuarine, Coastal and Shelf Science 63, 1–11.
DeGange, A.R. 1996. A conservation assessment for the marbled murrelet in southeast Alaska. U.S. Department
of Agriculture, Forest Service, Pacific Northwest Research Station, General Technical Report PNW-
GTR-388.
De Jong-Moreau, L., Casanova, B. & Casanova, J.-P. 2001. Detailed comparative morphology of the peri-oral
structures of the Mysidacea and Euphausiacea (Crustacea): an indication for the food preference.
DeLancey, L.B. 1987. The summer zooplankton of the surf zone at Folly Beach, South Carolina. Journal of
Coastal Research 3, 211–217.
Delgado, L., Guerao, G. & Ribera, C. 1997. Biology of the mysid Mesopodopsis slabberi (Van Beneden, 1861)
(Crustacea, Mysidacea) in a coastal laboon of the Ebro delta (NW Mediterranean). Hydrobiologia
357, 27–35.
127
PETER A. JUMARS
Deprez, T., Vanden Berghe, E. & Vincx, M. 2004. NeMys: a multidisciplinary biological information system.
In Proceedings ‘The Colour of Ocean Data’: International Symposium on Oceanographic Data and
Information Management with Special Attention to Biological Data, Brussels, Belgium, November
25–27, 2002, E. Vanden Berghe et al. (eds). International Oceanographic Commission (IOC) Workshop
Report 188, 57–63.
Deprez, T., Speybroeck, J., Steyaert, M. & Vincx, M. 2005. NeMys. World Wide Web electronic publication.
http://www. nemys.ugent.be, version and date accessed (7/2006).
De Robertis, A., Schell, C. & Jaffe, J.S. 2003. Acoustic observations of the swimming behavior of the
euphausiid Euphausia pacifica Hansen. ICES Journal of Marine Science 60, 885–898.
Dewicke, A., Cattrijsse, A., Mees, J. & Vincx, M. 2003. Spatial patterns of the hyperbenthos of subtidal
sandbanks in the southern North Sea. Journal of Sea Research 49, 27–45.
Dewicke, A., Rottiers, V., Mees, J. & Vincx, M. 2002. Evidence for an enriched hyperbenthic fauna in the
Frisian front (North Sea). Journal of Sea Research 47, 121–139.
Dieckmann, U., Law, R. & Metz, J.A.J. (eds) 2000. The Geometry of Ecological Interactions. Cambridge:
Cambridge University Press.
Divoky, G.J. 1978. Identification, documentation and delineation of coastal migratory bird habitat in Alaska.
2: Feeding habits of birds in the Beaufort Sea. In Environmental Assessment of the Alaskan Continental
Shelf. Annual Reports of Principal Investigators for the Year Ending March 1978. Volume 1 —
Mammals-Birds, J.H. Steele (ed.). Boulder, Colorado: U.S. Environmental Research Laboratories
Outer Continental Shelf Environmental Assessment Program, 549–569.
Donahoe, C.J. 1997. Age, growth, distribution and food habits of recently settled white seabass, Atractoscion
nobilis, off San Diego County, California. Fishery Bulletin, National Oceanic and Atmospheric
Administration of the United States 95, 709–721.
Drake, P., Arias, A.M., Baldó, F., Cuesta, J.A., Rodríguez, A., Silva-García, A., Sobrino, I., García-González,
D. & Fernández-Delgado, C. 2002. Spatial and temporal variation of the nekton and hyperbenthos
from a temparate European estuary with regulated freshwater inflow. Estuaries 25, 451–468.
Dunham, J.S. & Duffus, D.A. 2002. Diet of gray whales (Eschrichtius robustus) in Clayoquot Sound, British
Columbia, Canada. Marine Mammal Science 18, 419–437.
Dunne, J.A., Williams, R.J. & Martinez, N.D. 2004. Network structure and robustness of marine food webs.
Eleftheriou, A. & Holme, N.A. 1984. Macrofauna techniques. In Methods for the Study of Marine Benthos,
N.A.Holme & A.D. McIntyre (eds). Oxford: Blackwell Scientific, 140–216.
Elizalde, M., Dauvin, J.-C. & Sorbe J.-C. 1991. Les Mysidacès suprabenthiques de la marge sud du Canyon
du cap-ferret (Golfe de Gascogne): répartition bathymètrique et activité natatoire. Annales de l’Institut
océanographique 67, 129–144.
Emmerson, M. & Yearsley, J.M. 2004. Weak interactions, omnivory and emergent food-web properties.
Proceedings of the Royal Society of London B 271, 397–405.
Emsley, S.M., Tarling, G.A. & Burrows, M.T. 2005. The effect of vertical migration strategy on retention and
dispersion in the Irish Sea during spring-summer. Fisheries Oceanography 14, 161–174.
Enbysk, B.J. & Linger, F.I. 1966. Mysid statoliths in shelf sediments of northwest North America. Journal of
Sedimentary Petrology 36, 839–840.
Evans, M.S., Noguchi, G.E. & Rice, C.P. 1989. The biomagnification of polychlorinated biphenyls, toxaphene,
and DDT compounds in a Lake Michigan offshore food web. Archives of Environmental Contamination
and Toxicology 20, 87–83.
Feyrer, F., Herbold, B., Matern, S.A. & Moyle, P.B. 2003. Dietary shifts in a stressed fish assemblage: Conse-
quences of a bivalve invasion in the San Francisco Estuary. Environmental Biology of Fishes 67, 277–288.
Fleminger, A. & Clutter, R.I. 1965. Avoidance of towed nets by zooplankton. Limnology and Oceanography
10, 96–104.
Flinkman, J., Vuorinen, I. & Aro, E. 1992. Planktivorous Baltic herring (Clupea harengus) prey selectively on
reproducing copepods and cladocerans. Canadian Journal of Fisheries and Aquatic Sciences 49, 73–77.
Flynn, A.J. & Ritz, D.A. 1999. Effect of habitat complexity and predatory style on the capture success of fish
feeding on aggregated prey. Journal of the Marine Biological Association of the United Kingdom 79,
487–494.
128
Fogarty, M.J. & Murawski, S.A. 1998. Large-scale disturbance and the structure of marine systems: fishery
impacts on Georges Bank. Ecological Applications 8, Supplement: Ecosystem Management for Sus-
tainable Marine Fisheries, S6–S22.
Fosså, J.H. 1986. Aquarium observations on vertical zonation and bottom relationships of some deep-living
hyperbenthic mysids (Crustacea: Mysidacea). Ophelia 25, 107–117.
Fosså, J.H. & Brattegard, T. 1990. Bathymetric distribution of Mysidacea in fjords of western Norway. Marine
Foster, E.G., Ritz, D.A., Osborn, J.E. & Swadling, K.M. 2001. Schooling affects the feeding success of
Australian salmon (Arripis trutta) when preying on mysid swarms (Paramesopodopsis rufa). Journal
Frank, K.T., Petrie, B., Choi, J.S. & Leggett, W.C. 2005. Trophic cascades in a formerly cod-dominated
ecosystem. Science 308, 1621–1623.
Friesen, J.A., Mann, K.H. & Novitsky, J.A. 1986. Mysis digests cellulose in the absence of a gut flora. Canadian
Journal of Zoology 64, 442–446.
Froneman, P.W. 2001. Stable isotope (∂13C) composition of the food web of the temperate Kariega estuary
(Eastern Cape). African Journal of Aquatic Sciences 26, 49–56.
Fulton, R.S., III. 1982. Predatory feeding of two marine mysids. Marine Biology 72, 183–191.
Fulton, R.S., III. 1983. Interactive effects of temperature and predation on an estuarine zooplankton community.
Gal, G., Rudstam, L.G. & Greene, C.H. 1999. Acoustic characterization of Mysis relicta. Limnology and
Gama, A.M. da S., Montu, M.A. & de Gusmao, L.L.F. 2002. Ciclo de vida pós-marsupial e crescimento de
Metamysidopsis elongata atlantica (Crustacea, Mysidacea, Mysidae) em cultivo de laboratório. Iher-
ingia. Série Zoologia 92, 77–83.
Gatlin III, D.M. 1997. Nutrition and feeding of striped bass and hybrid striped bass. In Striped Bass and Other
Morone Culture, R.M. Harrell (ed.). Amsterdam: Elsevier, 235–252.
Genin, A. 2004. Bio-physical coupling in the formation of zooplankton and fish aggregations over abrupt
topographies. Journal of Marine Systems 50, 3–20.
Gibson, R.N. 2003. Go with the flow: tidal migration in marine animals. Hydrobiologia 503, 153–161.
Gibson, R.N. & Ezzi, I.A. 1980. The biology of the scaldfish, Arnoglossus laterna (Walbaum) on the west
coast of Scotland. Journal of Fish Biology 17, 565–575.
Goddard, S.M. & Forward, R.B. 1991. The role of the underwater polarized light pattern, in the sun compass
navigation of the grass shrimp, Paleomonetes vulgaris. Journal of Comparative Physiology A 169,
479–491.
González, L.A. 1974. Hallazgo de Neomysis americana Smith (1873) (Crustacea: Mysidacea) en el Río de
La Plata. Revista de Biología del Uruguay 11, 119–130.
Green, K. & Williams, R. 1986. Observations on food remains in faeces of elephant, leopard and crabeater
seals. Polar Biology 6, 43–45.
Greenlaw, C.F., Holliday, D.V. & McGehee, D.E. 2004. High-frequency scattering from saturated sand
sediments. Journal of the Acoustical Society of America 115, 2818–2823.
Grimm, V., Günther, C.-P., Dittmann, S. & Hildebrandt, H. 1999. Grid-based modelling of macrozoobenthos
in the intertidal of the Wadden Sea: Potentials and limitations. In The Wadden Sea, S. Dittmann (ed.).
Berlin: Springer, 207–226.
Grimm, V. & Railsback, S.F. 2005. Individual-Based Modeling in Ecology. Princeton, New Jersey: Princeton
University Press.
Grimm, V., Revilla, E., Berger, U., Jeltsch, F., Mooj, W.M., Railsback, S.F., Thulke, H.-H., Weiner, J., Wiegand,
T. & DeAngelis, D.L. 2005. Pattern-oriented modeling of agent-based complex systems: lessons from
ecology. Science 310, 987–991.
Hampel, H., Cattrijsse, A. & Vincx, M. 2003a. Habitat value of a developing estuarine brackish marsh for
fish and macrocrustaceans. ICES Journal of Marine Sciences 60, 278–289.
Hampel, H., Cattrijsse, A. & Vincx, M. 2003b. Tidal, diel and semi-lunar changes in the faunal assemblages
of an intertidal salt marsh creek. Estuarine, Coastal and Shelf Science 56, 795–805.
129
PETER A. JUMARS
Hamren, U. & Hansson, S. 1999. A mysid shrimp (Mysis mixta) is able to detect the odour of its predator
(Clupea harengus). Ophelia 51, 187–191.
Hanamura, Y. 1997. Review of the taxonomy and biogeography of shallow-water mysids of the genus Archaeo-
mysis (Crustacea: Mysidacea) in the North Pacific Ocean. Journal of Natural History 31, 669–711.
Hanamura, Y., Jo, S.-G. & Murano, M. 1996. A new species of Archaeomysis from coastal waters of Japan
(Mysidacea, Gastrosaccinae). Crustaceana 69, 553–566.
Hanamura, Y. & Matsuoka, M. 2003. Feeding habits of the sand shrimp, Crangon uritai Hayashi & Kim,
1999, in the central Seto Inland Sea, Japan. Crustaceana 76, 1017–1024.
Hanson, J.M. & Chouinard, G.A. 2002. Diet of Atlantic cod in the southern Gulf of St. Lawrence as an index
of ecosystem change, 1959–2000. Journal of Fisheries Biology 60, 902–922.
Hansson, S., Hobbie, J.E., Elmgren, R., Larsson, U., Fry, B. & Johansson, S. 1997. The stable nitrogen isotope
ratio as a marker of food-web interactions and fish migration. Ecology 78, 2249–2257.
Haury, L., Fey, C., Newland, C. & Genin, A. 2000. Zooplankton distribution around four eastern North Pacific
seamounts. Progress in Oceanography 45, 69–105.
Hays, G.C., Kennedy, H. & Frost, B.W. 2001. Individual variability in diel vertical migration of a marine
copepod: why some individuals remain at depth while others migrate. Limnology and Oceanography
46, 2050–2054.
Herman, S.S. 1963. Vertical migration of the opossum shrimp, Neomysis americana Smith. Limnology and
Hirota, Y., Koshiishi, Y. & Naganuma, N. 1990. Size of mysids eaten by juvenile flounder Paralichthys olivaceus
and diurnal change of its feeding activity. Bulletin of the Japanese Society of Fisheries 56, 201–206.
Hoffmeyer, M.S. 2004. Decadal change in zooplankton seasonal succession in the Bahía Blanca estuary, Argen-
tina, following introduction of two zooplankton species. Journal of Plankton Research 26, 181–189.
Holyoak, M. & Sachdev, S. 1998. Omnivory and the stability of simple food webs. Oecologia 117, 413–419.
Hong, S.Y. & Oh, C.W. 1989. Ecology of the sand shrimp, Crangon affinis in the Nakdong River estuary,
Korea. Bulletin of the Korean Fisheries Society 22, 351–362.
Hopkins, T.L. 1965. Mysid shrimp abundance in surface waters of Indian River Inlet, Delaware. Chesapeake
Science 6, 86–91).
Hostens, K. & Mees, J. 1999. The mysid-feeding guild of demersal fishes in the brackish zone of the
Westerschelde estuary. Journal of Fish Biology 55, 704–719.
Huang, M. 2004. Study on feeding habits and nutrient level of four cephalopod species from Taiwan Strait
and its adjacent areas. Journal of Oceanography in Taiwan Strait (Taiwan Haixia) 23, 331–339.
Hulbert, E.M. 1957. The distribution of Neomysis americana in the estuary of the Delaware River. Limnology
Hurlbert, S.N. 1971. The nonconcept of species diversity: a critique and alternative parameters. Ecology 52,
577–586.
Inoue, T., Suda, Y. & Sano, M. 2003. Food habits of fishes in the surf zone of a sandy beach at Sanrimatsubara,
Fukuoka Prefecture, Japan. Ichthyological Research 52, 9–14.
Jackson, C.J., Marcogliese, D.J. & Burt, M.D.B. 1997. Role of hyperbenthic crustaceans in the transmission
of marine helminth parasites. Canadian Journal of Fisheries and Aquatic Sciences 54, 815–820.
Janssen, J., Jones, W.R., Whang, A. & Oshel, P.E. 1995. Use of the lateral line in particulate feeding in the
dark by juvenile alewife (Alosa pseudoharengus). Canadian Journal of Fisheries and Aquatic Sciences
52, 353–363.
Jerling, H.L. & Wooldridge, T.H. 1995. Feeding of two mysid species on plankton in a temperate South
African estuary. Journal of Experimental Marine Biology and Ecology 188, 243–259.
Johannsson, O.E., Leggett, M.F., Rudstam, L.G., Servos, M.R., Mohammadian, M.A., Gal, G., Dermott, R.M.
& Hesslein, R.H. 2001. Diet of Mysis relicta in Lake Ontario as revealed by stable isotope and gut
content analysis. Canadian Journal of Fisheries and Aquatic Sciences 58, 1975–1986.
Johnson, W.S. & Allen, D.M. 2005. Zooplankton of the Atlantic and Gulf Coasts: A Guide to Their Identification
and Ecology. Baltimore, Maryland: Johns Hopkins University Press.
Johnston, N.M. & Ritz, D.A. 2001. Synchronous development and release of broods by the swarming mysids
Anisomysis mixta australis, Paramesopodopsis rufa and Tenagomysis tasmaniae (Mysidacea: Crusta-
cea). Marine Ecology Progress Series 223, 225–233.
130
Jumars, P.A. 1993. Concepts in Biological Oceanography. New York: Oxford University Press.
Jumars, P.A. 2000. Animal guts as ideal chemical reactors: maximizing absorption rates. American Naturalist
155, 527–543.
Jumars, P.A. & Nowell, A.R.M. 1984. Fluid and sediment dynamic effects on marine benthic community
structure. American Zoologist 24, 45–55.
Jumars, P.A., Self, R.F.L. & Nowell, A.R.M. 1981. A simple model of flow-organism-sediment interaction.
Marine Geology 42, 155–172.
Kaartvedt, S. 1985. Diel changes in small-scale vertical distribution of hyperbenthic mysids. Sarsia 70, 285–295.
Kaartvedt, S. 1989. Retention of vertically migrating suprabenthic mysids in fjords. Marine Ecology Progress
Series 57, 119–128.
Kaartvedt, S., Aksnes, D.L. & Aadnesen, A. 1988. Winter distribution of macroplankton and micronekton in
Masfjorden, western Norway. Marine Ecology Progress Series 45, 45–55.
Kaufmann, R.S., Smith, K.L., Jr., Baldwin, R., Glatts, R.C., Robison, B.H. & Reisenbichler, K.R. 1993.
Epipelagic communities in the northwestern Weddell Sea: results from acoustic, trawl and trapping
surveys. Antarctic Journal of the United States 28, 138–141.
Kemeny, J.G. & Snell, J.L. 1960. Finite Markov Chains. Princeton, New Jersey: Van Nostrand.
Kibirige, I., Perissinotto, R. & Nozais, C. 2003. Grazing rates and feeding preferences of the mysid shrimp
Gastrosaccus brevifissura in a temporarily open estuary in South Africa. Marine Ecology Progress
Series 251, 201–210.
Kim, S.L. & Oliver, J.S. 1989. Swarming benthic crustaceans in the Bering and Chuckchi seas and their
relation to geographic patterns in gray whale feeding. Canadian Journal of Zoology 67, 1531–1542.
Kimmerer, W.J. 2002. Physical, biological, and management responses to variable freshwater flow into the
San Francisco estuary. Estuaries 25, 1275–1290.
Kimmerer, W.J., Burau, J.R. & Bennett, W.A. 1998a. Persistence of tidally-oriented vertical migration by
zooplankton in a temperate estuary. Estuaries 25, 359–371.
Kimmerer, W.J., Burau, J.R. & Bennett, W.A. 1998b. Tidally-oriented vertical migration and position main-
tenance of zooplankton in a temperate estuary. Limnology and Oceanography 43, 1697–1709.
Kimmerer, W.J. & Orsi, J.J. 1996. Causes of long-term declines in zooplankton in the San Francisco Bay
estuary since 1987. In San Francisco Bay: The Ecosystem, J.T. Hollibaugh (ed.). Washington, D.C.:
American Association for the Advancement of Science, 403–424.
Köpcke, B. & Kausch, H. 1996. Distribution and variability in abundance of Neomysis integer and Mesopo-
dopsis slabberi (Mysidacea; Crustacea) in relation to environmental factors in the Elbe Estuary. Archiv
für Hydrobiologie. Supplementband. Untersuchungen des Elbe-Aestuars 110, 263–282.
Kotta, I. & Kotta, J. 2001a. Vertical migrations of mysids in the Gulf of Riga. Proceedings of the Estonian
Academy of Sciences. Biology, Ecology 50, 248–255.
Kotta, I. & Kotta, J. 2001b. Distribution of mysids on bank slopes in the Gulf of Riga. Proceedings of the
Estonian Academy of Sciences. Biology, Ecology 50, 14–21.
Kouassi, E., Pagano, M., Saint-Jean, L. & Sorbe, J.C. 2006. Diel vertical migrations and feeding behavior of
the mysid Rhopalophthalmus africana (Crustacea: Mysidacea) in a tropical lagoon (Ebrié, Côte
d’Ivoire). Estuarine, Coastal and Shelf Science 67, 355–368.
Kreeger, K.E., Kreeger, D.A., Langdon, C.J. & Lowry, R.R. 1991. The nutritional value of Artemia and
Tigriopus californicus (Baker) for two Pacific mysid species, Metamysidopsis elongata (Holmes) and
Mysidopsis intii (Holmquist). Journal of Experimental Marine Biology and Ecology 148, 147–158.
Kringel, K., Jumars, P.A. & Holliday, D.V. 2003. A shallow scattering layer: high-resolution acoustic analysis
of nocturnal vertical migration from the seabed. Limnology and Oceanography 48, 1223–1234.
Kunze, E., Dower, J.F., Beveridge, I., Dewey, R. & Bartlett, K.P. 2006. Observations of biologically generated
turbulence in a coastal inlet. Science 313, 1768–1770.
Lagardère, J.P. 1977. Recherches sur le régime alimentaire et le comportment prédateir dés decapodes
benthiques de la pente continentale de l’Atlantique nord orientale (Golfe de Gascogne et Maroc). In
Biology of Benthic Organisms, B.F. Keegan et al. (eds). Oxford: Pergamon Press, 397–408.
Lagardère, J.-P. & Nouvel, H. 1980. Les Mysidacés du talus continental du golfe de Gascogne. II. Familles
des Lophogastridae, Eucopiidae et Mysidae (Tribu des Erythropini exceptée). Bulletin du Muséum
national d’Histoire naturelle, 4ème série, section A, Zoologie, Biologie et Écologie animales 2, 375–412.
131
PETER A. JUMARS
Lake, S., Burton, H. & van den Hoff, J. 2003. Regional, temporal and time-scale spatial variation in Weddell
seal diet at four coastal locations in east Antarctica. Marine Ecology Progress Series 254, 293–305.
Lankford, T.E. & Targett, T.E. 1997. Selective predation by juvenile weakfish: post-consumptive constraints
on energy maximization and growth. Ecology 78, 1049–1061.
Lasenby, D.C. & Sherman, R.K. 1991. Design and evaluation of a bottom-closing net used to capture mysids
and other suprabenthic fauna. Canadian Journal of Zoology 69, 783–786.
Lasenby, D.C. & Shi, Y. 2004. Changes in the elemental composition of the stomach contents of the opossum
shrimp Mysis relicta during diel vertical migration. Canadian Journal of Zoology 82, 525–528.
Lawrie, S.M., Speirs, D.C., Raffaelli, D.G., Gurney, W.S.C., Paterson, D.M. & Ford, R. 1999. The swimming
behaviour and distribution of Neomysis integer in relation to tidal flow. Journal of Experimental
Marine Biology and Ecology 242, 95–106.
Lawson, G.L., Wiebe, P.H., Ashjian, C.J., Chu, D. & Stanton, T.K. 2006. Improved parameterization of
Antarctic krill target strength models. Journal of the Acoustical Society of America 119, 232–242.
Ledoyer, M. 1969. Écologie de la faune vagile des biotopes Méditerranéens accessible en scaphandre auto-
nome. V — Étude des phénomènes nycthéméraux. Les variations nycthémérales des populations
animals dans les biotopes. Tethys 1, 291–308.
Leising, A.W. 2001. Copepod foraging in patchy habitats and thin layers using a 2-D individual-based model.
Le Mao, P. 1985. Place de la seiche Sepia officinalis (mollusque, Cephalopoda) dans les chaines alimentaires
du golfe Normano-Breton. Cahiers de Biologie Marine 26, 331–340.
Linden, E. & Kuosa, H. 2004. Effects of grazing and excretion by pelagic mysids (Mysis spp.) on the size
structure and biomass of the phytoplankton community. Hydrobiologia 514, 73–78.
Linden, E., Lehtiniemi, M. & Viitasalo, M. 2003. Predator avoidance behaviour of Baltic littoral mysids
Neomysis integer and Praunus flexuosus. Marine Biology 143, 845–850.
Lindström M., Nilsson, H.L. & Meyer-Rochow, V.B. 1988. Recovery from light-induced sensitivity loss in
the eye of the crustacean Mysis relicta in relation to temperature: a study of ERG-determined V/log I
relationship and morphology at 4°C and 14°C. Zoological Science 5, 743–757.
Link, J.S. 2002. Does food web theory work for marine ecosystems? Marine Ecology Progress Series 230, 1–9.
Link, J.S. & Garrison, L.P. 2002. Trophic ecology of Atlantic cod Gadus morhua on the northeast U.S.
continental shelf. Marine Ecology Progress Series 227, 109–123.
Lock, K. & Mees, J. 1999. The winter hyperbenthos of the Ria Formosa — a lagoon in southern Portugal —
and adjacent waters. Cahiers de Biologie Marine 40, 47–56.
Longhurst, A.R., Bedo, A.W., Harrison, W.G., Head, E.J.H. & Sameoto, D.D. 1990. Vertical flux of respiratory
carbon by oceanic diel migrant biota. Deep-Sea Research Part A 37, 685–694.
Longhurst, A.R. & Harrison, W.G. 1988. Vertical nitrogen flux from the oceanic photic zone by diel migrant
zooplankton and nekton. Deep-Sea Research 35, 881–889.
Lowry, L.F. & Burns, J.J. 1980. Foods utilized by bowhead whales near Barter Island, Alaska, autumn 1979.
Marine Fisheries Review 42, 88–91.
Macquart-Moulin, C. & Patriti, G. 1993. Canyons sous marins et advection vers le talus continental du plancton
néretique. Oceanologica Acta 16, 179–189.
Macquart-Moulin, C. & Ribera Maycas, E. 1995. Inshore and offshore diel migrations in European bentho-
pelagic mysids, genera Gastrosaccus, Anchialina and Haplostylus (Crustacea, Mysidacea). Journal
of Plankton Research 17, 531–555.
Makris, N.C., Ratilal, P., Symonds, D.T., Jagannathan, S., Lee, S. & Nero, R.W. 2006. Fish population and
behavior revealed by instantaneous continental shelf-scale imaging. Science 311, 660–663.
Matsumoto, G.I. 1995. Observations on the anatomy and behaviour of the cubozoan Carybdea rastonii Haacke.
Marine and Freshwater Behaviour and Physiology 26, 139–148.
Mattila, J., Chaplin, G., Eilers, M.R., Heck, K.L., Jr., O’Neal, J.P. & Valentine, J.F. 1999. Spatial and diurnal
distribution of invertebrate and fish fauna of a Zostera marina bed and nearby unbegetated sediments
in Damariscotta River, Maine, U.S.A. Netherlands Journal of Sea Research 41, 321–332.
Mauchline, J. 1968. The biology of Erythrops serrata and E. elegans [Crustacea, Mysidacea]. Journal of the
Marine Biological Association of the United Kingdom 48, 455–464.
Mauchline, J. 1980. The biology of mysids and euphausiids. Advances in Marine Biology 18, 1–681.
132
Maurer, D. & Wigley, R.L. 1982. Distribution and ecology of mysids in Cape Cod Bay, Massachusetts, U.S.A.
Biological Bulletin (Woods Hole) 163, 405–530.
Mayer, L.M., Jumars, P.A., Taghon, G.L., Macko, S.A. & Trumbore, S. 1993. Low-density particles as potential
nitrogenous food sources for benthos. Journal of Marine Research 51, 373–389.
Mayer, L.M., Schick, L.L., Sawyer, T. & Plante, C.J. 1995. Bioavailable amino acids in sediments: a bio-
mimetic, kinetics-based approach. Limnology and Oceanography 40, 511–520.
McCann, K. & Hastings, A. 1997. Re-evaluating the omnivory-stability relationship in food webs. Proceedings
of the Royal Society of London B 264, 1249–1254.
McConnaughey, T. & McRoy, C.P. 1979. Food-web structure and the fractionation of carbon isotopes in the
Bering Sea. Marine Biology 53, 257–262.
Mees, J., Cattrijsse, A. & Hamerlynck, O. 1993. Distribution and abundance of shallow-water hyperbenthic
mysids (Crustacea, Mysidacea) and euphausiids (Crustacea, Euphausiacea in the Voordelta and the
Westerschelde, southwest Netherlands. Cahiers de Biologie Marine 34, 165–186.
Mees, J. & Hamerlynck, O. 1992. Spatial community structure of the winter hyperbenthos of the Schelde estuary,
the Netherlands, and the adjacent coastal waters. Netherlands Journal of Sea Research 29, 357–370.
Mees, J. & Jones, M.B. 1997. The hyperbenthos. Oceanography and Marine Biology: An Annual Review 35,
221–255.
Metillo, E.B. & Ritz, D.A. 2001. Laminarinase activity in three co-occurring mysids (Crustacea: Mysidacea)
from southeastern Tasmania, Australia. Philippine Scientist 38, 82–101.
Miller, C.B., Lynch, D.R., Carlotti, F., Gentleman, W. & Lewis, C.V.W. 1998. Coupling of an individual-based
population dynamic model of Calanus finmarchicus to a circulation model for the Georges Bank
region. Fisheries Oceanography 7, 219–234.
Moffat, A.M. & Jones, M.B. 1993. Correlation of the distribution of Mesopodopsis slabberi (Crustacea,
Mysidacea) with physico-chemical gradients in a partially-mixed estuary (Tamar, England). Nether-
lands Journal of Aquatic Ecology 27, 155–162.
Möllmann, C., Kornilovs, G., Fetter, M. & Köster, F.W. 2004. Feeding ecology of central Baltic Sea herring
and sprat. Journal of Fish Biology 65, 1563–1581.
Montoya, J.M., Pimm, S.L. & Solé, R.V. 2006. Ecological networks and their fragility. Nature 442, 259–264.
Moran, S. & Fishelson, L. 1971. Predation of a sand-dwelling mysid crustacean Gastrosaccus sanctus by
plover birds (Charadriidae). Marine Biology 9, 63–64.
Morris, D.W. 2005. Paradoxical avoidance of enriched habitats: have we failed to appreciate omnivores?
Ecology 86, 2568–2577.
Mouny, P., Dauvin, J.-C. & Zouhiri, S. 2000. Benthic boundary layer fauna from the Seine estuary (eastern
English Channel, France): spatial distribution and seasonal changes. Journal of the Marine Biological
Association of the United Kingdom 80, 959–968.
Mulkins, L.M., Jelinski, D.E., Karagatzides, J.D. & Carr, A. 2002. Carbon isotope composition of mysids at
a terrestrial-marine ecotone, Clayoquot Sound, British Columbia, Canada. Estuarine, Coastal and
Shelf Science 54, 669–675.
Navarro, J.C. & Villanueva, R. 2000. Lipid and fatty acid composition of early stages of cephalopods: an
approach to their lipid requirements. Aquaculture 183, 161–177.
Neff, J.M., 1997. Ecotoxicology of arsenic in the marine environment. Environmental Toxicology and Chem-
istry 16, 917–927.
Neil, D.M. 1975a. The control of eyestalk movements in the mysid shrimp Praunus flexuosus. Journal of
Experimental Biology 62, 487–504.
Neil, D.M. 1975b. The optokinetic responses of the mysid shrimp Praunus flexuosus. Journal of Experimental
Biology 62, 505–518.
Neil, D.M. 1975c. The mechanism of statocyst operation in the mysid shrimp Praunus flexuosus. Journal of
Experimental Biology 62, 685–700.
Nemerson, D.M. & Able, K.W. 2004. Spatial patterns in diet and distribution of juveniles of four fish species
in Delaware Bay marsh creeks: factors influencing fish abundance. Marine Ecology Progress Series
276, 249–262.
Oh, C.W., Hartnoll, R.G. & Nash, R.D.M. 2001. Feeding ecology of the common shrimp Crangon crangon
in Port Erin Bay, Isle of Man, Irish Sea. Marine Ecology Progress Series 214, 211–223.
133
PETER A. JUMARS
Ohtsuka, S., Inagaki, H., Onbe, T. & Yoon, Y.H. 1995. Direct observations of groups of mysids in shallow
coastal waters of western Japan and southern Korea. Marine Ecology Progress Series 123, 33–44.
Omori, M. 1978. Zooplankton fisheries of the world: a review. Marine Biology 48, 199–205.
Omori, M. & Hamner, W.M. 1982. Patchy distribution of zooplankton: behavior, population assessment and
sampling problems. Marine Biology 72, 193–200.
Orensanz, J.M., Schwindt, E., Pastorino, G., Bortolus, A., Casas, G., Darrigran, G., Elías, R., López Gappa,
J.J., Obenat, S., Pascual, M., Penchaszadeh, P., Piriz M.L., Scarabino, F., Spivak, E.D. & Vallarino,
E.A. 2002. No longer the pristine confines of the world ocean: a survey of exotic marine species in
the southwestern Atlantic. Biological Invasions 4, 115–143.
Orsi, J.J. 1986. Interaction between diel vertical migration of a mysidacean shrimp and two-layered estuarine
flow. Hydrobiologia 137, 79–87.
Orsi, J.J. & Mecum, W. 1986. Zooplankton distribution and abundance in the Sacramento-San Joaquin Delta
in relation to certain environmental factors. Estuaries 9, 326–339.
Papaconstantinou, C. & Caragitsou, E. 1987. The food of hake (Merluccius merluccius) in Greek seas. Vie et
Milieu 7, 77–83.
Patterson, H.M. 2004. Small-scale distributions and dynamics of the mysid prey of gray whales (Eschrictius
robustus) in Clayoquot Sound, British Columbia Canada. Ph.D. thesis, University of Victoria, British
Columbia, Canada.
Pearre, S., Jr. 2003. Eat and run? The hunger/satiation hypothesis in vertical migration: history, evidence and
consequences. Biological Reviews 78, 1–79.
Penry, D.L. & Jumars, P.A. 1987. Modeling animal guts as chemical reactors. American Naturalist 129, 69–96.
Perissinotto, R., Nozais, C., Kibirige, I. & Anandrai, A. 2003. Planktonic food webs and benthic-pelagic
coupling in three South African temporarily-open estuaries. Acta Oecologica 24(Suppl. 1), S307–S316.
Petryashev, V.V. 1992. Notes on mysid systematics (Crustacea, Mydidacea) of the Arctic and the north-western
Pacific Zoologichesky Zhurnal 71, 47–58.
Petryashev, V.V. 2002a. Leptostraca, Mysidacea, Isopoda, and Decapoda (Anomura) (Crustacea, Malacostraca)
of the Chukichi Sea and adjacent waters: hydrological regime and species composition. Russian
Journal of Marine Biology 28(Suppl. 1), S47–S55.
Petryashev, V.V. 2002b. Leptostraca, Mysidacea, Isopoda, and Decapoda (Anomura) (Crustacea, Malacostraca)
of the Chukichi Sea and adjacent waters: biogeography and fauna formation. Russian Journal of
Marine Biology 28(Suppl. 1), S56–S63.
Petryashev, V.V. 2005. Biogeographical division of the North Pacific sublittoral and upper bathyal zones by
the fauna of Mysidacea and Anomura (Crustacea). Russian Journal of Marine Biology 31(Suppl. 1),
S9–S26.
Pihl, L. & Rosenberg, R. 1984. Food selection and consumption of the shrimp Crangon crangon in some
shallow marine areas in western Sweden. Marine Ecology Progress Series 15, 159–168.
Plummer, K.M., Demartini, E.E. & Roberts, D.A. 1983. The feeding habits and distribution of juvenile-small
adult California halibut (Paralichthys californicus) in coastal waters off northern San Diego County.
Calilfornia Cooperative Fisheries Investigations Reports 24, 194–201.
Price, K.S., Jr. 1962. Biology of the sand shrimp, Crangon septemspinosa, in the shore zone of the Delaware
Bay region. Chesapeake Science 3, 244–255.
Price, W.W., Heard, R.W. & Stuck, L. 1994. Observations on the genus Mysidopsis Sars, 1864 with the
designation of a new genus, Americamysis, and the descriptions of Americamysis alleni and A. stucki
(Peracarida: Mysidacea: Mysidae), from the Gulf of Mexico. Proceedings of the Biological Society
of Washington 107, 680–698.
Quirt, J. & Lasenby, D. 2002. Cannibalism and ontogenetic changes in the response of the freshwater shrimp
Mysis relicta to chemical cues from conspecific predators. Canadian Journal of Zoology 80, 1022–1025.
Remerie, T., Bourgois, T., Peelaers, D., Vierstraete, A., Vanfleteren, J. & Vanreusel, A. 2006. Phylogeographic
patterns of the mysid Mesopodopsis slabberi (Crustacea, Mysida) in western Europe: evidence for
high molecular diversity and cryptic speciation. Marine Biology 149, 465–481.
Ribes, M., Coma, R., Zabala, M. & Gili, J.-M. 1996. Small-scale spatial heterogeneity and seasonal variation
in a population of a cave-dwelling Mediterranean mysid. Journal of Plankton Research 18, 659–671.
134
Rice, A.L. 1961. The response of certain mysids to changes in hydrostatic pressure. Journal of Experimental
Biology 38, 391–401.
Rice, A.L. 1964. Observations on the effects of changes of hydrostatic pressure on the behaviour of some
marine animals. Journal of the Marine Biological Association of the United Kingdom 44, 163–175.
Richoux, N.B., Deibel, D., Thompson, R.J. & Parrish, C.C. 2004. Seasonal changes in the lipids of Mysis
mixta (Mysidacea) from the hyperbenthos of a cold-ocean environment (Conception Bay, Newfound-
land). Canadian Journal of Fisheries and Aquatic Sciences 61, 1940–1953.
Richoux, N.B., Deibel, D., Thompson, R.J. & Parrish, C.C. 2005. Seasonal and developmental variation in
the fatty acid composition of Mysis mixta (Mysidacea) and Acanthostepheia malmgreni (Amphipoda)
from the hyperbenthos of a cold-ocean environment (Conception Bay, Newfoundland). Journal of
Plankton Research 27, 719–733.
Ritz, D.A. 1994. Social aggregation in pelagic invertebrates. Advances in Marine Biology 30, 155–216.
Roast, S.D., Widdows, J. & Jones, M.B. 1998. The position maintenance behaviour of Neomysis integer
(Peracarida: Mysidacea) in response to current velocity, substratum and salinity. Journal of Experi-
mental Marine Biology and Ecology 220, 25–45.
Roast, S.D., Widdows, J., Pope, N. & Jones, M.B. 2004. Sediment-biota interactions: mysid feeding activity
enhances water turbidity and sediment erodability. Marine Ecology Progress Series 281, 145–154.
Rooney, N., McCann, K., Gellner, G. & Moore, J.C. 2006. Structural asymmetry and the stability of diverse
food webs. Nature 442, 265–269.
Rudstam, L.G., Danielsson, K., Hansson, S. & Johansson, S. 1989. Diel vertical migration and feeding patterns
of Mysis mixta (Crustacea, Mysidacea) in the Baltic Sea. Marine Biology 101, 43–52.
Saigusa, M., 2001. Daily rhythms of emergence of small invertebrates inhabiting shallow subtidal zones: a
comparative investigation at four locations in Japan. Ecological Research 16, 1–28.
Sameoto, D. 2001. Decadal changes in phytoplankton color index and selected calanoid copepods in continuous
plankton recorder data from the Scotian Shelf. Canadian Journal of Fisheries and Aquatic Sciences
58, 749–761.
Sanger, G.A. 1987. Winter diets of common murres and marbled murrelets in Kachemak Bay, Alaska. The
Condor 89, 426–430.
San Vicente, C. & Munilla, T. 2000. Misidáceaos suprabentónicos de las laysa catalanas (Mediterráneo
nordoccidentale). Orsis 15, 45–55.
Sardiña, P. & Lopez Cazorla, A.C. 2005a. Feeding habits of the juvenile striped weakfish, Cynoscion guatucupa
Cuvier 1830, in Bahá Blanca estuary (Argentina): seasonal and ontogenetic changes. Hydrobiologia
532, 23–38.
Sardiña, P. & Lopez Cazorla, A.C. 2005b. Trophic ecology of the whitemouth croaker, Micropogonias furnieri
(Pisces: Sciaenidae), in south-western Atlantic waters. Journal of the Marine Biological Association
of the United Kingdom 85, 405–413.
Sawamto, S. 1987. Mysids collected in Amitori Bay, Iriomote Island, Ryukyu Islands. Bulletin of Marine
Science 41, 642 (abstract only).
Schiariti, A., Berasategui, A.D., Giberto, D.A., Guerrero, R.A., Acha, E.M. & Mianzan, H.W. 2006. Living
in the front: Neomysis americana (Mysidacea) in the Rio de la Plata estuary, Argentina-Uruguay.
Schindler, D.E. & Scheuerell, M.D. 2002. Habitat coupling in lake ecosystems. Oikos 98, 177–189.
Schlacher, T.A. & Wooldridge, T.H. 1994. Tidal influence on distribution and behaviour of the estuarine
opossum shrimp Gastrosaccus brevifissura. In Changes in Fluxes in Estuaries: Implications from
Science to Management, K.D. Dyer & R.J. Orth (eds). Fredensborg, Denmark: Olsen & Olsen,
307–312.
Shields, J.D. 1994. The parasitic dinoflagellates of marine crustaceans. Annual Reviw of Fish Diseases 4,
241–271.
Siebeck, O. 1968a. ‘Uferflucht’ und optische Orientierung pelagischer Crustaceen. Archiv für Hydrobiologie
(Suppl.) 35, 1–118.
Siebeck, O. 1968b. Spatial orientation of planktonic crustaceans 2. The swimming behaviour in a vertical plane.
Verhandlungen Internationale Vereinigung für theoretische und angewandte Limnologie 17, 841–847.
135
PETER A. JUMARS
Siegfried, C.A. 1982. Trophic relations of Crangon franciscorum Stimpson and Palaemon macrodactylus
Rathbun: predation on the opossum shrimp, Neomysis mercedis Holmes. Hydrobiologia 89, 129–139.
Sitts, R.M. & Knight, A.W. 1979. Predation by the estuarine shrimps Crangon franciscorum Stimpson and
Palaemon macrodactylus Rathbun. Biological Bulletin (Woods Hole) 156, 356–368.
Smedbol, R.K. & Stephenson, R. 2001. The importance of managing within-species diversity in cod and
herring fisheries of the north-western Atlantic. Journal of Fish Biology 59(Suppl. A), 109–128.
Smith, S.I. 1873. Report upon the invertebrate animals of Vineyard Sound and the adjacent waters, with an
account of the physical characters of the region. Report of the U.S. Fish Commission for 1871–1872,
295–747 [description of Neomysis americana on pp. 552–553].
Smith, S.L., Pieper, R.E., Moore, M.V., Rudstam, L.G., Greene, C.H., Zamon, J.E. & Williamson, C.E. 1992.
Acoustic techniques for the in situ observation of zooplankton. Archiv für Hydrobiologie — Advances
in Limnology 36, 23–43.
Song, K.-H. & Breslin, V.T. 1999. Accumulation and transport of sediment metals by the vertically migrating
opossum shrimp, Mysis relicta. Journal of Great Lakes Research 25, 429–442.
Sorbe, J.C. 1991. Biología del misidaceo suprabentónico Schistomysis ornata (Sars, 1864) en la platforma
continental aquitana (suroeste de Francia). Actas V Simposium Ibérica Estudios Bentos Marinas 1,
273–298.
Speirs, D., Lawrie, S., Raffaelli, D., Gurney, W. & Emes, C. 2002. Why do shallow-water predators migrate?
Strategic models and empirical evidence from an estuarine mysid. Journal of Experimental Marine
Spencer, C.N., Potter, D.S., Bukantis, R.T. & Standford, J.A. 1999. Impact of predation by Mysis relicta on
zooplankton in Flathead Lake, Montana, U.S.A. Journal of Plankton Research 21, 51–64.
Stelle, L.L. 2002. Behavioral ecology of summer resident gray whales (Eschrichtius robustus) feeding on
mysids in British Columbia, Canada. Ph.D. thesis, University of California, Los Angeles.
St.-Hilaire, A., Courtenay, S.C., Dupont, F. & Boghen, A.D. 2002. Diet of white perch (Morone americana)
in the Richibucto estuary, New Brunswick. Northeastern Naturalist 9, 303–316.
Stone, H.H. & Daborn, G.R. 1987. Diet of alewives, Alosa pseudoharengus and blueback herring, A. aestivalis
(Pisces: Clupeidae) in Minas Basin, Nova Scotia, a turbid, macrotidal estuary. Environmental Biology
of Fishes 19, 55–67.
Suh, H.-L., Jo, S.-G. & Kim, K.-Y. 1995. Diel horizontal migration of the two mysids Archaeomysis kokuboi
and Acanthomysis sp. in the sandy shore surf zone of Yongil Bay, eastern Korea. Journal of the
Oceanological Society of Korea 30, 523–528.
Sutherland, D.L. & Closs, G.P. 2001. Diel patterns of mysid drift (Crustacea: Mysidacea) in the Taieri River
estuary, New Zealand. New Zealand Journal of Marine and Freshwater Research 35, 197–200.
Takahashi, K. 2004. Feeding ecology of mysids in freshwater and coastal marine habitats: a review. Bulletin
of the Plankton Society of Japan 51, 46–72.
Takahashi, K., Hirose, T., Azuma, N. & Kawaguchi, K. 2004. Diel and intraspecific variation in vulnerability
of the beach mysid, Archaeomysis kokuboi Ii, 1964, to fish predators. Crustaceana 77, 717–728.
Takahashi, K., Hirose, T. & Kawaguchi, K. 1999. The importance of intertidal sand-burrowing peracarid
crustaceans as prey for fish in the surf-zone of a sandy beach in Otsuchi Bay, northeastern Japan.
Fisheries Science 65, 856–864.
Takahashi, K. & Kawaguchi, K. 1995. Inter- and intraspecific zonation in three species of sand-burrowing
mysids, Archaeomysis kukuboi, A. grebnitzkii and Iiella ohshimai, in Otsuchi Bay, northeastern Japan.
Takahashi, K. & Kawaguchi, K. 1997. Diel and tidal migrations of the sand-burrowing mysids, Archaeomysis
kokuboi, A. japonica and Iiella ohshimai, in Otsuchi Bay, northeastern Japan. Marine Ecology Progress
Series 148, 95–107.
Takeuchi, T., Park, G.S., Seikai, T. & Yokoyama, M. 2001. Taurine content in Japanese flounder Paralichthys
olivaceus T. & S. and red sea bream Pagrus major T. & S. during the period of seed production.
Aquaculture Research 32(Suppl. 1), 244–248.
Tanaka, Y., Yamaguchi, H., Gwak, W.-S., Tominaga, O., Tsusaki, T. & Tanaka, M. 2005. Influence of mass
release of hatchery-reared Japanese flounder on the feeding and growth of wild juveniles in a nursery
ground in the Japan Sea. Journal of Experimental Marine Biology and Ecology 314, 137–147.
136
Tararam, A.S., Wakabara, Y. & Flynn, M.N. 1996. Suprabenthic community of the Cananeia lagoon estuarine
region, southeastern Brazil. Cahiers de Biologie Marine 37, 295–308.
Tattersall, W.M. 1938. The seasonal occurrence of mysids off Plymouth. Journal of the Marine Biological
Taylor, L.H., Shellito, S.M., Abello, H.U. & Jumars, P.A. 2005. Tidally cued emergence events in a strongly
tidal estuary. Estuaries 28, 500–509.
Thorne, R.E. 1968. Diel variations in distribution and feeding behavior of Mysidacea in Puget Sound. M.S.
thesis, University of Washington, Seattle.
Tominaga, O., Uno, N. & Seikai, T. 2003. Influence of diet shift from formulated feed to live mysids on the
carbon and nitrogen stable isotope ratio (∂13C and ∂15N). Aquaculture 218, 265–276.
Tudela, S. & Palomera, I. 1997. Trophic ecology of the European anchovy Engraulis encrasicolus in the
Catalan Sea (northwest Mediterranean). Marine Ecology Progress Series 160, 121–134.
Twining, B.S., Gilbert, J.J. & Fisher, N.S. 2000. Evidence of homing behavior in the coral reef mysid Mysidium
gracile. Limnology and Oceanography 45, 1845–1849.
Vadeboncoeur, Y., McCann, K.S., Vander Zanden, M.J. & Rasmussen, J.B. 2005. Effects of multi-chain
omnivory on the strength of trophic control in lakes. Ecosystems 8, 682–693.
Vallet, C. & Dauvin, J.-C. 1998. Composition and diversity of the benthic boundary layer macrofauna from
the English Channel. Journal of the Marine Biological Association of the United Kingdom 78, 387–409.
Vallet, C. & Dauvin, J.-C. 2001. Biomass changes and bentho-pelagic transfers in the English Channel. Journal
of Plankton Research 23, 903–922.
Vallet, C., Zouhiri, S., Dauvin, J.-C. & Wang, Z. 1995. Variations nycthémérales de l’abondance de la faune
démersal en Manche. Journal de Recherches Océanographiques 20, 94–102.
van der Baan, S.M. & Holthuis, L.B. 1971. Seasonal occurrence of Mysidacea in the surface plankton of the
southern North Sea near the ‘Texel’ lightship. Netherlands Journal of Sea Research 5, 227–239.
Viherluoto, M., Kuosa, H., Flinkman, J. & Viitasalo, M. 2000. Food utilisation of pelagic mysids, Mysis mixta
and M. relicta, during their growing season in the northern Baltic Sea. Marine Biology 136, 553–559.
Viherluoto, M. & Viitasalo, M. 2001. Effect of light on the feeding rates of pelagic and littoral mysid shrimps:
a trade-off between feeding success and predation avoidance. Journal of Experimental Marine Biology
and Ecology 261, 237–244.
Viitasalo, M., Flinkman, J. & Viherluoto, M. 2001. Zooplanktivory in the Baltic Sea: a comparison of prey
selectivity by Clupea harengus and Mysis mixta, with reference to prey escape reactions. Marine
Vinogradov, M.E. 1962. Feeding of the deep-sea zooplankton. Rapports et Procès-Verbaux des Réunions du
Conseil International pour l’Exploration de la Mer 153, 114–119.
Visser, A.W. & Kiørboe, T. 2006. Plankton motility patterns and encounter rates. Oecologia 148, 538–546.
Voicu, G. 1981. Upper Miocene and recent mysid statoliths in central and eastern Paratethys. Micropaleon-
tology 27, 227–247.
Wahle, R.A. 1985. The feeding ecology of Crangon franciscorum and Crangon nigricauda in San Francisco
Bay, California. Journal of Crustacean Biology 5, 311–326.
Wang, Z. & Dauvin, J.-C. 1994. The suprabenthic fauna of the infralittoral fine sand community from the
Bay of Seine (eastern English Channel) — composition, swimming activity and diurnal variation.
Cahiers de Biologie Marine 35, 135–155.
Warkentine, B.E. & Rachlin, J.W. 1989. Winter offshore diet of the Atlantic silverside, Menidia menidia.
Copeia 1989, 195–198.
Waterman, T.H. 2005. Reviving a neglected celestial underwater polarization compass for aquatic animals.
Biological Reviews 81, 1–5.
Webb, P., Perissinotto, R. & Wooldridge, T.H. 1987. Feeding of Mesopodopsis slabberi (Crustacea, Mysidacea)
on naturally occurring phytoplankton. Marine Ecology Progress Series 38, 115–123.
Webb, P., Perissinotto, R. & Wooldridge, T.H. 1988. Diet and feeding of Gastrosaccus psammodytes (Crus-
tacea, Mysidacea) with special reference to the surf zone diatom Anaulus birostratus. Marine Ecology
Webb, P. & Wooldridge, T.H. 1990. Diel horizontal migration of Mesopodopsis slabberi (Crustacea: Mysida-
cea) in Algoa Bay, southern Africa. Marine Ecology Progress Series 62, 73–77.
137
PETER A. JUMARS
Whiteley, G.C., Jr. 1948. The distribution of larger planktonic Crustacea on Georges Bank. Ecological
Wicklum, D. 1999. Variation in horizontal zooplankton abundance in mountain lakes: shore avoidance or fish
predation? Journal of Plankton Research 21, 1957–1975.
Wigley, R.L. 1967. Comparative efficiencies of Van Veen and Smith-McIntyre grab samplers as revealed by
motion pictures. Ecology 48, 168–169.
Wigley, R.L. & Burns, B.R. 1971. Distribution and biology of mysids (Crustacea, Mysidacea) from the Atlantic
coast of the United States in the NMFS Woods Hole collection. Fishery Bulletin, National Oceanic
and Atmospheric Administration of the United States 69, 717–746.
Wigley, R.L. & Theroux, R.B. 1981. Atlantic continental shelf and slope of the United States — macrobenthic
invertebrate fauna of the Middle Atlantic Bight Region — faunal composition and quantitative
distribution. Geological Survey Professional Paper 529-N, N1–N198.
Wilhelm, F.M., Hamann, J. & Burns, C.W. 2002. Mysid predation on amphipods and Daphnia in a shallow
coastal lake: prey selection and effects of macrophytes. Canadian Journal of Fisheries and Aquatic
Sciences 59, 1901–1907.
Williams, A.B. 1972. A 10-year study of meroplankton in North Carolina estuaries: mysid shrimps. Chesapeake
Science 13, 254–262.
Williams, A.B., Bowman, T.E. & Damkaer, D.M. 1974. Distribution, variation and supplemental description
of the opossum shrimp Neomysis americana (Crustacea: Mysidacea). Fishery Bulletin, National
Oceanic and Atmospheric Administration of the United States 72, 835–842.
Wing, B.L. & Barr, N. 1977. Midwater invertebrates from the southeastern Chukchi Sea: species and abundance
in catches incidental to midwater trawling survey of fishes, September–October, 1970. National
Oceanic and Atmospheric Administration of the United States, Technical Report NMFS SSRF-710,
1–43.
Winkler, G., Dodson, J.J., Bertrand, N., Thivierge, D. & Vincent, W.F. 2003. Trophic coupling across the St.
Lawrence River estuarine transition zone. Marine Ecology Progress Series 251, 59–73.
Winkler, G. & Greve, W. 2004. Trophodynamics of two interacting species of estuarine mysids, Praunus
flexuosus and Neomysis integer, and their predation on the calanoid copepod Eurytemora affinis.
Journal of Experimental Biology and Ecology 308, 127–146.
Wittman, K.J. 1977. Modifications of association and swarming in North Adriatic Mysidacea in relation to
habitat and interacting species. In Biology of Benthic Organisms, B.F. Keegan et al. (eds). Oxford:
Pergamon Press, 605–612.
Wittman, K.J. 2001. Centennial changes in the near-shore mysid fauna of the Gulf of Naples (Mediterranean
Sea), with description of Heteromysis riedli sp. n. (Crustacea, Mysidacea). Pubblicazioni della Stazi-
one Zoologica di Napoli I: Marine Ecology 22, 85–109.
Wittman, K.J. & Ariani, A.P. 1996. Some aspects of fluorite and vaterite precipitation in marine environments.
Pubblicazioni della Stazione Zoologica di Napoli I: Marine Ecology 17, 213–219.
Yamamoto, M., Makino, H., Kobayashi, J.-I. & Tominaga, O. 2004. Food organisms and feeding habits of
larval and juvenile Japanese flounder Paralichthys olivaceus at Ohama Beach in Hiuchi-Nada, the
central Seto Inland Sea, Japan. Fisheries Science 70, 1098–1105.
Yamamoto, M. & Tominaga, O. 2005. Feeding ecology of dominant demersal fish species Favonigobius
gymnauchen, Repomucenus spp. and Tarphops oligolepis at a sandy beach where larval Japanese
flounder settle in the Seto Inland Sea, Japan. Fisheries Science 71, 1332–1340.
Zagursky, G. & Feller, R.J. 1985. Macrophyte detritus in the winter diet of the estuarine mysid Neomysis
americana. Estuaries 8, 355–362.
Zhivotovsky, L.A., Bergman, A. & Feldman, M.W. 1996. A model of individual adaptive behavior in a
fluctuating environment. In Adaptive Individuals in Evolving Populations, R.K. Belew & M. Mitchell
(eds). Santa Fe, New Mexico: Santa Fe Institute, 131–153 (Santa Fe Institute Studies in the Sciences
of Complexity 26).
Zouhiri, S. & Dauvin, J.-C. 1996. Diel changes of the Benthic Boundary Layer macrofauna over coarse sand
sediment in the western English Channel. Oceanologica Acta 19, 141–153.
Zouhiri, S., Vallet, C., Mouny, P. & Dauvin, J.-C. 1998. Spatial distribution of suprabenthic mysids from the
English Channel. Journal of the Marine Biological Association of the United Kingdom 78, 1181–1202.
138

Taylor & Francis
USE OF DIVERSITY ESTIMATIONS IN THE STUDY

OF SEDIMENTARY BENTHIC COMMUNITIES
ROBERT S. CARNEY
Department of Oceanography and Coastal Sciences, Louisiana State University,
Baton Rouge, Louisiana, U.S.A. 70803
E-mail: rcarne1@lsu.edu
Abstract The soft-bottom benthos covers most of the sea floor. Measurement and analysis of the
species richness of these habitats are increasingly needed for studies of community regulation and
for providing scientific criteria for the conservation of the ocean bottom at all depths. Diversity
measures provide an evolving suite of tools that allow benthic ecologists to meet both basic and
applied needs. While species diversity is now considered a fundamental aspect of communities and
ecosystems, the measurement of benthic diversity did not become commonplace until the late 1960s.
Prior to that communities were characterised by representative species with the implicit assumption
that minor species components did not warrant detailed analysis. Use of diversity measures in
benthic ecology has largely parallelled studies in other ecosystems with an emphasis upon measures
that are informative when applied to large amounts of data with high species numbers. Non-parametric
indices such as Simpson’s and Shannon’s are widely used along with simple species richness. Log-
series and log-normal distributions have been advocated as general neutral models but receive less
use. Current research is especially focused upon extrapolation of unsampled species richness and
diversity relationships across spatial scales. Major contributions from benthic ecology include the
rarefaction of samples to a uniform size, the development of indices that include phylogenetic
relationships in diversity estimation and the extrapolation of full species richness from observed
values. In meeting scientific and societal needs, benthic ecologists must apply methods that are
insightful yet can be simply explained within the resource-policy arena.
Introduction
Justification
Estimation of diversity has become an integral part of benthic ecology. There is so much recent
literature and software available that review may seem unneeded. Benthic ecology is, however,
now experiencing a change in the ways that species data are accessed and analytical results used
that is both scientific and societal in origin. Both origins require that concepts and estimation of
diversity be reconsidered. The greatest scientific change is the increasing accessibility of survey
data through open Internet databases. This allows the search for geographic and temporal patterns
not anticipated in the original study designs and a search across multiple studies by experts in
analysis and theory who may be largely unfamiliar with benthic ecology and the taxonomy of
benthic organisms. The second change is societal in the sense that international regulatory policies
increasingly mandate the preservation of biological diversity in both marine and terrestrial systems.
Benthic ecologists must provide regulators with estimates of diversity that can be explained and
defended if these estimates are to serve agencies as the basis for conservation decisions. Thus, the
intent of this review is to provide users of databases an explanation of what benthic ecologists have
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ROBERT S. CARNEY
found and provide benthic ecologists a guide to the changes associated with the shift in terminology
from diversity to biodiversity.
Contraction of the term biological diversity to biodiversity seems to have originated within the
U.S. government environmental management structure and was then progressively used by those
ecologists especially interested in conservation biology (Harper & Hawksworth 1994). Along with
development of conservation biology, biodiversity began to encompass a much broader concept
than species diversity alone and now may be considered a distinct concept or suite of concepts
(Hamilton 2005). One marine definition of biodiversity included the variety of genomes, species
and ecosystems occurring in a defined region (National Research Council 1995) and followed a
similar combination of genetic and ecological perspectives used by Norse and his colleagues (Norse
et al. 1986).
The official definition of biodiversity as contained in Article 2 of the Convention on Biological
Diversity included “variability among living organisms from all sources … within species, between
species, and of ecosystems” (United Nations Conference on Environment and Development, 1992).
The view adopted in this review is that biodiversity is largely a policy term rather than scientific
and its use should be avoided. Efforts to better define biodiversity from a scientific standpoint are
needed and reflect conservation biologists’ duty to provide objective tools to managers faced with
mandates to preserve biodiversity in marine as well as terrestrial systems (Lubchenco et al. 2003).
Presently, however, policy usage of ‘biodiversity’ carries with it many assumptions that have not
been proven scientifically such as a link between diversity of ecosystem health (Norse 1993) and
ecosystem stability.
Notable efforts in ecology to provide management tools were the adoption in benthic ecology
of taxonomic indices that weight diversity by phylogenetic differences (Warwick & Clarke 2001)
and the search for indicator species to be used in place of more comprehensive diversity assessment.
As discussed in the historical review, selection of indicator species bears a strong similarity to the
selection of characteristic species during the decades of benthic ecology research prior to any
interest in the diversity of bottom communities.
‘Biodiversity informatics’ is the term applied to the growing development and use of databases
for diversity studies and is very broadly defined to include biogeography and certain aspects of
systematics. Progress and challenges for systems that will provide marine data have been outlined
by Costello & Berghe (2006). There is already progress for deep-sea studies starting with data
compiled by many French cruises (Fabri et al. 2006) and by many studies conducted in shallow
European seas (Costello et al. 2006). Initially, these marine databases can most confidently be used
for determining geographic and bathymetric ranges of individual species. As problems of incon-
sistent and incorrect taxonomy are solved, however, the datasets will be extremely useful for
estimating benthic diversity over a wide range of scales.
Structure of the review

This review takes a broad historical perspective to examine how benthic ecology has treated diversity
from approximately 1870 until the present time with special attention to soft bottoms. Benthic
ecologists carried out surveys as early as the 1900s that were similar to the projects of today, but
lacked both the modern concepts of diversity and the computational tools to compute indices.
However, there are strong similarities between the struggle of early benthic ecologists to simplify
discussion of species-rich systems and the search of contemporary conservation biologists for
indicator taxa that can be used in the estimation of overall community diversity (Pearson 1994).
The mathematics of diversity estimation are treated herein only in sufficient detail to indicate
what benthic ecologists do and have done with respect to concepts and data analysis. Only those
approaches widely used in or originating in benthic ecology are considered. Texts by Hayek &
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USE OF DIVERSITY ESTIMATIONS IN THE STUDY OF SEDIMENTARY BENTHIC COMMUNITIES
Buzas (1997) and Magurran (2004) do an excellent job of focusing upon the major concepts and
methods. The former text has somewhat greater mathematical detail, while the latter text provides
more information about concept development. Even these recent books are quickly outdated.
Methods, concepts and large-scale patterns of diversity with respect to mud bottoms have been
considered in highly informative reviews of Gray (2000, 2001, 2002). The information presented
herein is intended to compliment these works by taking a broader historical perspective and tracing
the use of analytical tools more than by discussing details of many individual results. Unfortunately,
all reviews must choose to omit something. The two serious omissions here are (1) the use of
evenness measures to compliment diversity and (2) the effect of pollution stress on benthic diversity.
Both topics warrant separate treatment in the future. In concluding, recommendations are made as
to a future course in benthic ecology that will allow both a better understanding of diversity and
an ability to provide managers with useful information.
Basics
To avoid contributing to additional confusion, it is necessary to state the concept of diversity used
in this review. According to a simple view of systems ecology, there are three types of information
about a benthic community (Figure 1). First, an ‘inventory’ is a list of all species and their
abundance. Second, a set of interactions among the component species is often represented by a
matrix. Third, a set of relationships exists between the fauna and the physical environment. Sam-
pling, identification and enumeration produce the inventory. Determination of fauna-environment
relationships can be made through sampling designs that capture variation in sediment type, salinity,
temperature, and so on. Assessment of species interactions is the most difficult information to
obtain. Certainly, soft-bottom communities are impractical locations to determine the population
interaction parameters required by theoretical community matrices (Levins 1968). In some cases,
however, associations such as dependence on biogenic structure are obvious and a variety of tools
can be used to determine at least a trophic position. The assumption is that the abundance of each
species in the inventory can be explained to some extent by the interactions among species and the
interactions with the environment.
Of these three sets of information, diversity is an attribute of the inventory (Peet 1974). When
given a mathematical definition, diversity should afford a parsimonious means of comparing the
inventories of different systems. The underlying assumption is that differences in diversity reflect
differences in species interactions. Common questions in benthic ecology have been directed to
whether ubiquitous gradients of diversity exist with depth, with latitude and with anthropogenic
stress. In each case, diversity is a convenient indicator of ecosystem differences.
Terminology varies greatly in the larger ecological literature, but most authors take the position
advocated by Hill (1973) and Hurlbert (1971). Measures of species diversity (the variety of the
inventory) are based on two simple attributes: the number of species (species richness) and the pro-
portional abundances. An effective means of describing the variability of proportional abundance is
evenness (i.e.,departure from equal proportions). Using these two attributes, indices can be calcu-
lated and used as an overall measure of heterogeneity (Magurran 2004).
A somewhat unsettling aspect about species diversity is that all species are treated equally,
making no use of additional knowledge about biotic or abiotic interactions and life histories. Failure
to treat some species as more important would seem to make a traditional species diversity measure
poorly suited to be used for conservation decisions about which communities should be afforded
special protections. A partial solution is seen in a recent development in benthic ecology, use of
indices of taxonomic distinctness (Warwick & Clarke 2001). Still an attribute of the inventory,
these indices make use of additional information about taxonomic position of the component
species. The adoption of these indices marks a major change in benthic community analysis.
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ROBERT S. CARNEY
Species-by-sample quantitative data
Sample 1
Sample 2
Sample 3
Sample 4
Sample 5
Sample 6
Sample 7
Sample 8
Sample 9
Sample 10
Sample k
Species 1 x1,1 x1,2 x1,3 x1,4 x1,5 x1,6 x1,7 x1,8 x1,9 x1,10 … x1,k
…
…
Species i xi,1 xi,2 xi,3 xi,4 xi,5 xi,6 xi,7 xi,8 xi,9 xi,10 … xi,k
Speciesenvironment factor interactions
Temperature
Sediment
Factor m
Factor 5
Factor 6
Factor 7
Factor 8
Salinity
Depth
Species 1 ρ1,1 ρ1,2 ρ1,3 ρ1,4 ρ1,5 ρ1,6 ρ1,7 ρ1,8 … ρ1,m
Speciespair relation matrix Species 2 ρ2,1 ρ2,2 ρ2,3 ρ2,4 ρ2,5 ρ2,6 ρ2,7 ρ2,8 … ρ2,m
Species 1
Species 2
Species 3
Species 4
Species 5
Species 6
Species i
Species 1 1 α1,2 α1,3 α1,4 α1,5 α1,6 … α1,j
…
…
…
…
Species 2 α2,1 1 α2,3 α2,4 α2,5 α2,6 … α2,j Species i xi,1 xi,2 xi,3 xi,4 xi,5 xi,6 xi,7 xi,8 … xi,m
Species 3 α3,1 α3,2 1 α3,4 α3,5 α3,6 … α3,j
Species 4 α4,1 α4,2 α4,3 1 α4,5 α4,6 … α4,j
Species 5 α5,1 α5,2 α5,3 α5,4 1 α5,6 … α5,j
Species 6 α6,1 α6,2 α6,3 α6,4 α6,5 1 … α6,j
…
…
…
…
…
…
…
Species i αi,1 αi,2 αi,3 αi,4 αi,5 αi,6 … 1
Figure 1 Basic nature of soft-bottom benthic survey data. Ecology theory takes the position that population
levels of individual species in a community are influenced by interactions with the environment, including
resource utilisation, and pairwise relationships among species. In application, benthic surveys produce quan-
titative species-by-sample data according to designs that nest replicates with stations within larger ocean areas.
Interactions of species with the environment are often expressed as correlation coefficients and are limited to
the few factors included in the sampling design. An actual matrix of the relationships among pairs of species
is rarely known, but statistical associations are sometimes developed as substitutes from the species-sample
data. Traditionally, species diversity has been seen as a property of the species-by-sample data alone, ignoring
the other two types of data.
History
From Forbes zones to Petersen communities
When benthic studies from the late 1800s through the mid-1900s are reviewed a peculiar situation
emerges about use of species diversity. Early hints of interest in diversity existed prior to the advent
142
of community ecology, but then there was surprisingly little interest during early formative years of
community ecology. Finally tremendous new interest began in the 1950s as niche theory and easy
computation facilitated inquiry. Certainly, benthic surveys produced inventories in which a few
species were common and many more rare, but comments as to this fact are largely absent from
about 1900 to 1960. With so much emphasis upon diversity today, it is informative to consider a
historical period of very active benthic surveying when the concept seems to have been missing or
unimportant.
Estimation of species diversity is now associated with quantitative benthic sampling. Toward
the end of the 1800s, seafloor studies began the transition from the description of faunal zones
based upon qualitative trawl and dredge sampling (Forbes 1859, Mills 1978, Carney 2005) to more
quantitative grab and core surveys. Interest in species diversity during qualitative sampling can be
seen from the criticism of the CHALLENGER Expedition (1872–1876) by Anton Stuxburg (1883).
Stuxburg complained about the lack of synthesis in the largely taxonomic works and specifically
suggested that the number of species and the proportions of each be presented trawl by trawl.
Possibly accepting these suggestions, the summary of the expedition issued 12 yr later carefully
noted that deep samples contained a greater variety of megafauna species that showed lower
numerical dominance than shallow samples in spite of the numerically smaller catch (Murray 1895).
No explanation of this higher deep diversity was presented, and the observation was largely forgotten,
possibly due to the much greater emphasis upon quantitative shallow water studies that soon followed.
Contemporary surveys of soft bottom benthic communities are distinguished by a strong
emphasis on numerical analysis of truly quantitative samples of the fauna in a known volume of
sediment lying under a similarly known area of the sea floor. The origin of this type of surveying
is generally attributed to the work of pioneering fisheries ecologist, C.G.J. Petersen (Petersen 1918),
The method was developed during the course of ecologically comprehensive fish stock assessment
begun in the late 1880s.
Petersen-type surveys producing species inventories were widely adopted. Local surveys were
conducted around Great Britain at such locations as in the vicinity of the Plymouth Marine
Laboratory (Ford 1923, Smith 1932) and Scotland (Stephen 1928, 1934, Clark & Milne 1955).
Numerous surveys took place along other west European coasts such as off Iceland and in the
Mediterranean. By the 1900s larger scale surveys were conducted in the English Channel (Holme
1966). In North America, Allee (1923) surveyed the benthos in the vicinity of Woods Hole. Possibly
most influential were benthic surveys in Puget Sound on the Pacific coast by Shelford (1935) who
was a strong proponent of the super-organism view of community structure and function. Similar
surveys were spread across the Arctic from the 1920s onward, and were summarised in English
by Zenkevitch (1963). The techniques were also adopted along the Japanese coast in the 1930s
and 1940s by Miyada (cited by Thorson 1957).
These many Petersen-type surveys were all quite similar although sampling gear and sediment
processing evolved over the course of the studies (Spärck 1935, Thorson 1955). The general trend
was towards larger areas of sampling and more reliable penetration of the bottom. Statistical
analyses were minimal, and results were often presented as a map of both faunal assemblages and
oceanographic conditions. Assemblages were inventoried in detail, then described and named on
the basis of the two characteristic species. Graphics were used to portray the relative abundance
of dominant species.
Diversity, as an aspect of the species inventories, was neither discussed nor analyzed in studies
into the 1960s. This was despite the availability of useful indices since the 1940s, and their
widespread use terrestrially for both plant and insect surveys. In addition these early workers
considered themselves to be studying communities as interacting systems. However, hints exist that
questions about species diversity were beginning to be formulated. In the survey by Smith (1932)
of the Eddystone grounds species richness was presented with singletons and more abundant species
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ROBERT S. CARNEY
carefully noted. Possibly reflecting growing ideas and better calculators, more sophisticated analyses
began to appear such as the dispersion of species across samples (Clarke & Milne 1955). By the
time of the English Channel survey (Holme 1966), the Petersen tradition of naming assemblages
after two characteristic species had been dropped due to the finding that species composition varied
greatly within such assemblages.
The surprisingly little interest in species diversity or in any related characterisation of species
inventories probably had several causes. The three most likely are a lack of practical utility, a lack
of relevant concepts, and a lack of computational tools. With respect to utility, many of these benthic
surveys were associated with fisheries studies making community productivity the parameter of
interest. The apparent lack of ideas about species diversity may be related to the immaturity of the
community concept. In the early 1900s, mapping of communities and characterisation of their
component species was the major activity, and not a careful investigation of community structure
and function that might be implied from the species inventory.
Jones (1950) reviewed the status of benthic studies in the context of community theory and
concluded that many workers accepted the idea that they were studying integrated systems in which
biological interactions were important. Few, however, seemed to fully embrace the idea that benthic
communities were superorganisms passing through biologically controlled successive states until
a certain climax was reached. Indeed, the distribution of benthic assemblages was always explained
in terms of control by physical conditions such as depth, sediment type, salinity, etc. One notable
exception was Shelford, who was one of the framers of the climax community and biome concepts
(Clements & Shelford 1939). He divided the oceans into a series of biomes largely associated with
depth and geographic position without reference to species richness. Another ecology pioneer was
Allee (1934), a strong proponent of benthic communities functioning as superorganisms, tracing
the idea back to Verrill.
At the end of Petersen era

In 1957, the state of knowledge about benthic ecology was compiled by international experts in a
twenty nine-chapter memoir and published by the Committee on Marine Ecology and Paleoecology
of the Geological Society of America (Hedgepeth 1957). Of particular relevance to the concept of
diversity was the paper on bottom communities by Thorson (1957). This paper clearly marks a
transition from the era of naming communities to one of discussing diversity patterns. The level
mud bottom was correctly seen as one of the largest, and apparently homogenous, environments
on Earth. Due to the strong dependence upon oceanographic conditions, bottom communities with
similar taxonomic structure should be found over very large areas. These parallel communities
were viewed as having relatively minor differences around the world.
More importantly, Thorson compiled species richness data on selected taxa and found an
increase from pole to tropics for epifauna and no gradient for infauna. Strongly influenced by
physiological explanations, the increase was attributed to greater thermal stability in the tropics. A
different view of benthic community stability emerged based upon ‘Thorson’s Rule’, a generalisa-
tion about increased occurrence of pelagic larvae in the tropics seen as having many exceptions
but some general validity (Laptikhovsky 2006). It was then suggested that the tropical benthos
would show greater spatial and temporal variation in species composition because of a large
variation in survival to settlement in the plankton. Higher latitudes should have a more stable
community structure due to the prevalence of direct development.
The strong emphasis on parsimoniously characterising multispecies communities in a manner
suitable for mapping without actual mathematical analyses lead early benthic ecologists to depend
on nomenclature, or the naming of communities. A reading of the very detailed “ideal rules” of
Thorson (1957) indicates how subjective the process actually was. Recommendations on how to
144
select characteristic species would be of only historical interest if a similar need did not exist today
to simply describe benthic communities for conservation planning. Later in this review it will be
shown that naming Petersen communities is similar to picking indicator species and assigning
greater importance to some species than others.
The primary task of naming communities was to identify within the collected fauna those
species that are ‘characteristic’ of the community. The five rules of Thorson paraphrased here were.
First, more than one such species should be selected. Second, short life-span species should be
avoided because their numbers fluctuate too much to be consistently characteristic. Third, highly
mobile animals and predators should be avoided as being be too transient. Fourth, characteristic
species should be big enough and abundant enough to be immediately conspicuous and have good
identification traits without consultation with a specialist. Fifth, biomass and/or density can be used
an indicator of abundance as long as they are not misleading due to large brood sizes or very large
specimens.
Even within the mundane task of picking names for communities, an interest in diversity can
be seen. Thorson divided the species inventory into four categories or orders based on abundance
and fidelity of association with a particular community. A first-order characteristic species should
be conspicuous, found throughout the range of the community in at least 50% of the samples, and
at least 5% of the biomass and restricted to that community. A second-order characteristic species
should have a similar frequency of occurrence and biomass dominance, but limited to only portions
of the range. A third-order species would be found in other communities as well as in at least 70%
of the units and at least 10% biomass. A fourth-order of ‘associated animals or influents’ would
be in at least 25% of the units and as much as 2% of the biomass but of little diagnostic value due
to a wide distribution crossing other communities.
Beginning of a new era

While formative elements of modern ecological theory may be found in many lines of early
population research, ecological questions about niche filling, resource utilisation, and competitive
exclusion were first expressed by G.E. Hutchinson and his students and colleagues in the 1960s
(Maurer 1999). The “diversity of a species inventory” was modelled as a balance achieved through
competition, resource specialisation, habitat complexity, resource availability, and history (Mac-
Arthur 1972). The details of community structure and function were being examined with mathe-
matical tools, and species diversity was a parameter of great interest.
The transition to the new view is most evident in a series of benthic studies begun in shallow
estuaries (Sanders 1960) and then extended to abyssal depths (Sanders et al. 1965). Initially, com-
munities were still named on the basis of characteristic species such as the Nephthys incisa – Nucula
proxima community, and diversity indices were not calculated (Sanders 1960). By 1965, descriptive
habitat names were used in place of characteristic species, and new diversity tools were proposed.
There was obvious interest in species richness and proportions, the large number of rarer species,
and the quantitative analysis of recurrent groups using trellis diagrams. Sanders’ benchmark com-
parative study of marine benthic diversity (Sanders 1968) marked the beginning of an adoption of
niche theory and analytical methods by benthic ecologists worldwide that persists to this day.
This comprehensive paper by Sanders made four major contributions. First, it objectively
examined the use of several diversity measures, and found that the information-based Shannon’s
index was adequate, but species richness was preferred. Secondly, rarefaction, a procedure for
estimating species richness in computationally reduced samples was presented to reduce the effect
of sample size. Third, Thorson’s infauna versus epifauna latitude gradients were challenged and
regional oceanographic conditions considered to be of greater importance in controlling diversity
highs and lows. Fourth, the high diversity of deep-sea macrofauna was noted for the first time since
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ROBERT S. CARNEY
the CHALLENGER Expedition and proposed as a general ocean feature. A stability-time hypothesis
was proposed as a general model for all benthic environments. In this explanation physical instability
was predicted to cause low diversity and biological accommodation would cause high diversity where
physical conditions were stable.
Sanders was extremely careful about making a distinction between measurements of diversity
that are reflective of species number (species diversity) and those reflective of proportional abun-
dance (dominance diversity). Although categorising several indices as being of one or the other
category, Sanders employed his own method of using species number per sample size for species
diversity. His method of calculating dominance diversity was to first plot a species accumulation
curve for each sample. He then compared that curve at reduced sample sizes (arrived at by
rarefaction) with a baseline curve representing maximum equitability with all species having the
same proportional abundance. Unfortunately, full details of the method were omitted.
Sanders proceeded to examine the behaviour of species diversity versus dominance diversity
in eight benthic habitats reducing the sample size artificially through rarefaction. A graphical means
was employed to track changes in rank of diversity as samples were rarefied. The ranks determined
by species number were found to be fairly consistent upon rarefaction, while ranks determined by
dominance were very inconsistent. He concluded that species number was the more conservative
measure of diversity while dominance was more variable due to the physical environment.
Influx of indices
The 1960s and 1970s saw a rapid adoption of diversity measures and multivariate approaches to
the analysis of benthic data. This adoption was due to a more fully developed niche theory, a better
access to computers, and a dissatisfaction with the subjectivity of Petersen-like community descrip-
tion (Lie personal communication). The origins of the indices, however, preceded adoption by
benthic ecologists by a decade or more.
The inventories, lists and counts of species, found in benthic or any other type of survey
sampling are categorical data in which individual specimens are assigned to a species category.
Linguists also deal with categorical data, and pioneers like Zipf (1935) and Yule (1944) developed
quantitative methods of comparing texts. They counted the frequency of words in various texts,
ordered those frequencies by rank and noted recurrent curves reflecting the fact that a few words
were very common and many rare. At roughly the same time period, R.A. Fisher (Fisher et al.
1943) proposed the use of a logarithmic series for examination of species categorical data. Influ-
enced by the linguistic indices, Simpson (1949) proposed use of a ‘concentration’ index, and
Shannon (1948) developed Information Theory that would be embraced by ecologists following a
suggestion by Margalef (1958).
The literature on how diversity should be measured continues to grow rapidly. Works in general
ecology published in the 1960s through 1980s tend to fall into a either a category dealing with
niche-theory models or a more practical category trying to improve the utility of indices. Benthic
studies of diversity fit into both categories, but place emphasis on practical aspects. The emphasis
on practical aspects stems from the increased number of surveys required to address environmental
problems. Both theoretical and practical works are now on an upsurge. Increased theoretical interest
has been generated by the proposal by Hubbell (2001) of the “unified theory of biodiversity and
biogeography” and by multinational interest in the preservation of the European coastal seas. The
‘unified theory’ has inspired considerable controversy (Whitfield 2002) and renewed examination
of diversity models (Pueyo 2006). Preservation of the coastal seas of many European nations
requires standardised measures of diversity that are both scientifically meaningful and useful for
policy and management decisions.
146
Compared with terrestrial studies, the use of diversity measures by benthic ecologists has been
relatively conservative in terms of restricting the types of indices proposed and applied. This can
be attributed to the nature of benthic survey data, that is, a collection of many thousands of
individuals and several hundred species. The taxonomy for many of the benthic groups is poorly
developed and often in need of revision. Many species are rare. Compounding these problems,
attempts at larger-scale syntheses are hindered by inconsistent sampling methods and great natural
variation in sample size. Therefore, benthic ecologists have always needed measures that were
robust when data were not ideal and which simplified the task of interpretation. Most studies have
made use of just a few diversity measures based either upon fitting abundance distribution models
or calculating an index. Most of these measures were well described by Gray (1981a) in benthic
terms. In the context of this review, use of a distribution means fitting and calculation of the
parameters that generate the distribution. Use of an index means the combining of two or more
characteristics of species-abundance distributions to produce a single value on a scale that allows
comparison among communities. Indices make no assumptions about the underlying distribution,
but carry with them implicit definitions of diversity. Use of distributions always allows for signif-
icance testing. For all common indices statistical properties have been developed and formal testing
is also possible.
Traditional approaches
Diversity measures are so widely applied and improved measures are so actively sought that a
division into traditional versus newer approaches is somewhat artificial. Old approaches are con-
stantly being reconsidered. That acknowledged, there are some approaches that have been in use
a long time and have been quite extensively discussed. These shall be presented first. Then some
of the more recent developments are considered.
Log-series and log-normal abundance distributions
From a statistical perspective the most parsimonious means of describing diversity and conducting
rigorous comparisons among communities is to first identify the underlying species abundance
distribution, and fit the model and estimate the parameters that characterise the distribution. Several
such distributions have been used in diversity studies (Hayek & Buzas 1997, Magurran 2004), but
the two oldest have had the greatest usage in benthic ecology. These are the log-series (Fisher et al.
1943) and log-normal (Preston 1948) distributions. The finding that either one or the other of these
distributions fitted a wide variety of terrestrial and marine data was once considered to reflect
profound aspects about ecosystem structure (Odum et al. 1960), and that studies of pattern alone
could definitively identify the causative processes. It has now been realised, however, that such
distributions may simply reflect the outcome of many complex processes, especially when there
are a large number of species and individuals are present (May 1975, Pueyo 2006). Indeed,
information on species abundance alone is insufficient to select among alternate ecological theories
of causation (McGill 2003). Many different processes can generate the same distribution.
Explanation of distributions, the process of fitting, and the determination of parameters is
substantially more complex than a discussion of diversity indices. Hayek & Buzas (1997) provide
an excellent detailed account, but these authors are strong advocates for the wide application of
the log-series. The log-series can be characterised using only a single parameter Fisher’s α.
Computing α requires an interactive computation. When data actually fit the log-series, α is
approximately the number of species represented by a single specimen (singletons).
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ROBERT S. CARNEY
An especially successful use of the log-series in benthic systems was an application to archived
foraminiferan data from five coastal regions ranging from the Arctic into the Caribbean (Buzas &
Culver 1989). Fisher’s α provided a highly useful measure of diversity and indicated a strong
geographic trend with the highest diversities in the tropical Caribbean and lowest in the Arctic. An
unusual aspect of that analysis was that log-series rarefaction was used (Hayek & Buzas 1997) to
produce equivalency, and that occurrence among samples was used as a measure of abundance
rather than counts within a sample.
The log-normal refers to abundances that are normally distributed about a mean once the data
have been log transformed. As for any normal distribution, it is characterised by two parameters —
mean and variance, which can be used as indicators of diversity. The log-normal has a rich history
of usage in ecology since first recognised as a widespread pattern (Preston 1948). An early
application in benthic ecology was the re-examination by Gage & Tett (1973) of benthic data from
two lochs that had been previously analyzed using rarefacted species richness (Gage 1972). The
log-normal distribution was fitted, and resulting means and variances used to search for patterns
associated with the salinity differences of two lochs, salinity gradient within each loch, and sediment
type. In the authors’ opinion, the two log-normal parameters provided a more informative picture
than rarefacted species richness. The actual goodness of fit, however, can be questioned since single-
tons were excluded before analysis. The complete data may have been better fitted with the log-series.
The most extensive use of the log-normal distribution in benthic ecology can be found in the
studies of John Gray and his colleagues. Gray (1981a) noted that benthic assemblages containing
many singletons generally fit the log-series distribution, but the common assemblage in which most
species were represented by a few individuals fit the log-normal. The log-normal distribution has
proven useful in identifying pollution impacts on benthic diversity (Gray 1981b, 1983, 1985). The
log-normal has been proposed as a neutral model for soft bottom macrofauna assemblages in the
sense that it is the expected outcome of certain ubiquitous processes of immigration, emigration,
and resources partitioning (Ugland & Gray 1982, 1983). In a renewed discussion about the genera-
tion of species abundance patterns by neutral models, the appropriateness of the log-normal has
been criticised (Williamson & Gaston 2005). Grey et al. (2006a), however, considered both a
terrestrial and a marine system, and argue that many systems may be effectively modelled as
compound log-normals in which two or more distributions are mixed. Ecologically, it seems quite
feasible that benthic samples will include several suites of species for which the abundances reflect
separate and distinct histories. Additional investigation is required.
Species richness and its rarefaction
Species richness is defined as the number of species in the samples of interest. Those samples may
represent replicates from a single location or from larger spatial scales. The notation and nomen-
clature of Gray (2000) serves to avoid confusion with other symbols and ambiguity as to scale.
‘SR’ denotes species richness with subscripts applied to indicate spatial extent. It is the most easily
explained of all measures of diversity, and for a large segment of the concerned community it is
synonymous with biodiversity. In his classification of indices (Hill 1973), “SR” is viewed as giving
equal weight to species of any abundance since it ignores those abundances completely. Recognising
that SR is a function of sample size N, SR is often normalised through division by N or area
sampled. Additionally, relationships of SR with sample size and abundance can be examined through
regression with the slope of a regression serving as a index of diversity. These approaches are well
covered by Hayek & Buzas (1997) and Magurran (2004). Species richness is often plotted against
sampling effort represented by counts, number of samples, or area sampled as an indication of the
completeness of the species inventory. In the case of a complete inventory, the curve becomes
asymptotic.
148
An important advancement in examination of species-effort and species-area curves was the

generation of multiple plots of subsets of the data selected at random by computer (Colwell 2005).
This produced both means and variances rather than single points along a curve. Renewed interest
in such relationships is based upon the potential to extrapolate species richness beyond the actual
level of sampling to be discussed below (Colwell & Coddington 1994, Ugland et al. 2003). Ana-
lytical approaches have, however, replaced randomisation.
The best application of species richness as a diversity measure is in a situation where the biota
has been fully inventoried with all species collected and recognised. This seldom if ever occurs in
benthic ecology. Rare species go unsampled due to insufficient sample size, and fine distinctions
between similar-appearing species can be easily overlooked. The problem of taxonomic error is
quite hard to overcome, but an adjustment can be made for differences in SR arising from unequal
sample size. Rarefaction originated in benthic studies (Sanders 1968) and has been widely adopted
throughout ecology. Its purpose is to reduce multiple samples to a common N, and then estimate
the number of species that should be present. Sanders also noted that the curves generated by
rarefaction proceeding through a range of N’s could also be used to rank samples by diversity.
Sanders use of rarefaction was not intended as a rigorous exercise in probabilities, and is best
considered as an instruction set for reducing sample size. Hurlbert (1971) and Simberloff (1972)
recognised the estimation of SR as a problem that could be solved by making use of the hyper-
geometric distribution and introduced the term expected species E(Sn) where Sn denotes species
richness at the reduced sample size. Rarefaction is no longer limited just to estimating SR, but to
other diversity measures as well using the hypergeometric and other distributions. Hayek & Buzas
(1997) compared four rarefaction methods using tree survey data. The hypergeometric produced
the best results, but Sander’s methods still proved both useful and simple.
Simpson’s λ
Simpson’s λ is an index based upon the probabilities encountered when comparing any two
individuals in a set of species. These probabilities are estimated from the proportional abundance
of each species in an assemblage. When two individuals are drawn, they may either be the same
or a different species. All possible outcomes can be displayed as a square matrix (Figure 2). The
diagonal of the matrix contains the probability of all possible ways in which the individuals drawn
are in the same species. The values above and below the diagonal are all the possible ways that
dissimilar species could be drawn. Since the order in which the species is found is unimportant,
the probabilities above and below the diagonal are equal. The sum of all terms in the matrix are
equal to one since no other combinations for two individuals exist. Simpson’s λ is the sum of all
the elements on the diagonal where S equals the number of species (Equation 1).
Simpson’s index λ= ∑p
i =1
2
(1)
Simpson’s λ was proposed (Simpson 1949) as a measure of the concentration of the classification
of individuals into species. The index has great conceptual appeal since it is the likelihood that two
individuals drawn at random without replacement from a community or sample of a community
belong to the same species. Terminology varies somewhat among users with Simpson’s D usually
refering to the form 1 – λ which has the preferred property of increasing with greater diversity.
The index can also be expressed expressed as 1/ λ , 1 – λ , and ln(λ) (Magurran 2004). The form
1 – λ is the probability of drawing two individuals that are not the same species (Equation 2). The
double summation indicates that summing of the elements excludes the diagonal. Only half the matrix
149
ROBERT S. CARNEY
Species abundance plots
0.25
Proportional abundance
Proportional abundances
0.20 for index calculation
Species 1
Species 2
Species 3
Species 4
Species 5
Species 6
Species j
0.15
Log-series?
0.10
Species 1 p1
2 p1 p2 p1 p3 p1 p4 p1 p5 p1 p6 … p1 pj
0.05
Species 2 p2 p1 p22 p2 p3 p2 p4 p2 p5 p2 p6 … p2 pj
.....
1 10 20 300 S
Rank of abundance
… … … … … … … …
50
Species i pi p1 pi p2 pi p3 pi p4 pi p5 pi pj … pi2
40
Species number
Species × species probability of all pairs

30
Log-normal?
20
10
1
1 10 50 100
Specimen count (logn scale)
Figure 2 Distributions or calculation of indices. Two common means of plotting species abundance, rank of
abundance versus proportion of sample and number of species versus number of individuals have led to the
suggestion that either the log-series or log-normal distribution could parsimoniously describe the data. Alter-
nately indices can be calculated, most often using the proportion of abundance. Proportions provide an estimate
of the probabilities that pairs of individuals drawn from the data will be the same (values on the diagonal) or
different (off the diagonal) species.
is summed according to this notation, requiring that the result be multiplied by two to get the actual
probability. It has come into renewed application as a component of taxonomic distinctiveness
discussed below. The index is sometimes referred to as the Gini-Simpson index in recognition of
development of the same function by the economist C. Gini in 1912 (Gorelick 2006).
S −1 S
1− λ = 2∑∑ p p
j=2 i< j
i j (2)
In his classification of indices (Hill 1973), λ gives greatest weight to abundant species. This
behaviour has reduced its popularity in benthic ecology and other fields that commonly encounter
numerous species with low abundances. For example, the index goes unmentioned in Gray (1981a).
The emphasis on abundant species is a property of squaring the proportions. Proportions are always
equal to or less than one. When proportions are squared the product is an even smaller fraction.
150
Thus, if the most dominant species in a sample has a p = 0.30, p2 = 0.090. A species with half that
abundance, p = 0.15 will contribute p2 = 0.023 to the summed index, or only a fourth as much.
The positive side of λ’s insensitivity to rare species is that it is minimally influenced by sample
size because abundant species are usually sampled with low effort. Therefore, λ produces relatively
consistent rankings of the least to the most diverse assemblages Lande et al. (2000). It has also
been effectively used to show latitude gradients in intertidal mudflats (Attrill et al. 2001), and
warrants greater consideration for similar comparisons across multiple studies. The abundant species
that most influence λ are most likely to be the best surveyed and most consistently and correctly
identified
Information theory and Shannon’s H′
Shannon’s index is the summation of plog(p) for all S species (Equations 3a,b).
Shannon’s H′ = − ∑ p log p
i =1
i i (3a)
H′ = − ∑ p ln( p )
i =1
i i (3b)
Unlike the conceptually simple Simpson’s λ , Shannon’s H′ is based on the more abstract field of
information theory and systems entropy (Shannon 1948). The formula appeared much earlier in
Boltzman’s 1872 work in entropy (Gorelick 2006) and simultaneously in the cybernetics work of
Weiner (1948). The index is sometimes termed the Shannon-Weiner index or incorrectly Shannon-
Weaver due to citation confusion (Magurran 2004). In spite of unclear conceptual relevance to
ecology, it continues in widespread use due to its mathematical properties and history of application.
Information theory provides a means to quantify the complexity of information that can be used
in the design of communication systems (Shannon 1948). It originated during World War II as a
tool for assuring the successful transmission and reception of encoded messages through noisy
radio channels. Its use in systems ecology for the quantification of diversity was first advocated by
Margalef (1958) on the basis of an analogy between transmission systems and temporal changes
in ecosystems. Very simplified, temporal changes are like a noisy channel between the structure of
an ecosystem at one time and another time. Pielou (1966) was very influential in the adoption of
information diversity measures, but specifically rejected the underlying analogy (Pielou 1969).
Margalef (1995) continued to advocate the utility of the analogy.
H′ is a fundamentally different way of envisioning diversity, and is related to the complexity
of the task of sorting the specimens into correct species groups through a series of decisions.
Compared to other measures of diversity, information has two very important distinguishing features
associated with the summed term plog(p), most often calculated as the natural logarithm pln(p).
First, pln(p) is modal reaching a maximum of 0.3679 for a proportion of p = 0.3678. Higher and
lower proportions contribute less to the summed index. Illustrating this point with an unlikely
assemblage of two species with proportions of 0.999994 and 0.000006, both the very common and
the very rare would contribute roughly equally to H′, approximately 0.000006 for both. Second,
H′ increases linearly with geometric increase of species richness under conditions of full evenness.
For example, if there are three assemblages with 10, 20 and 40 equally abundant species, the
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ROBERT S. CARNEY
respective H′ will be 2.303, 2.996 and 3.689. The increment in H′ is a consistent 0.693 even though
the species richness doubles. Depending upon one’s concept of diversity, these are either good or
bad properties.
Use of information theory for diversity quantification in benthic studies had been initiated by
the late 1960s (Lie and Kelly 1970, Lie 1974). Its popularity in benthic studies can be seen from the
fact that it is the only diversity index presented in Gray’s (1981a) succinct text on benthic ecology.
This popularity continues to the present (Gobin and Warwick 2006, Warwick et al. 2006). A variety
of diversity specialists have found the properties of H′ poorly suited for specific tasks (May 1975,
Lande 1996), and Magurran (2004) attributes its continued use largely to tradition. H′ does, however,
have properties very useful in diversity analysis. Specifically, it supports additive formulations of
diversity across scales from sample to large area (Lande 1996, Veech et al. 2002), and identification
of the underlying distribution of proportions can be made through examining the changes in species
richness, Equitability, and H′ during subsampling of data (SHE analysis, Hayek & Buzas 1997).
Newer developments
New developments in the measure of benthic diversity still fall into both theoretical and practical
categories, although there is greater merger of the two than previously. When sedimentary habitats
are sampled, the process of developing high quality species count data is far more time and effort
consuming than parallel activities such as chemical and granulometric analyses. Once the benthic
data are available, confusion can exist in explaining the data analyses applied. In the real situation
when both time and money are critical, there is a great emphasis upon doing things more expediently
and providing more informative results. The use of surrogates to estimate diversity is an approach
seeking to reduce effort. The use of new taxonomic diversities is an effort to improve results. A
bit closer to theory are attempts to extrapolate from small samples to larger areas, and to gain
knowledge over larger spatial scales by compiling local studies.
Surrogates
The intent of the surrogate approach is to replace the hard and expensive task of compiling a
multispecies inventory with an easier and less costly survey of indicator species, coarser taxonomic
level, or restricted size class. Proof that any of these surrogates are useful rests in demonstrating
that they allow for an accurate estimation of the diversity of unsampled species. Weaker proof is
that the surrogate produces a similar diversity ranking of assemblages as that obtained by more
comprehensive methods. Benthic ecologists are largely accepting that such approaches might work
if proven, since surrogacy is almost always applied to some extent. Benthic systems like most
others are complex, and benthic ecologists have traditionally met the need to adopt a practical focus
by dealing with a restricted size range or taxonomic category.
The concept of an indicator or surrogate for full diversity measurement has been widely
examined for terrestrial systems (Gaston & Williams 1993, Williams & Gaston 1994, Anderson
1995, Andelman & Fagan 2000). Unfortunately, approaches from the use of single species to more
inclusive groupings have shown little utility for reflecting diversity of the unsurveyed species
(Eduardo & Grelle 2002, MacNally et al. 2002, Su et al. 2004).
When the criteria for indicator species developed by Pearson and associates (Pearson 1994)
for conservation biology are critically examined, they seem intended to produce simple descriptors
of a community rather than to serve as a surrogate for diversity. Indeed, they are similar to rules
for identifying characteristic species in Petersen-type communities (Thorson 1957). Indicator cri-
teria can be rephrased as:
152
1. The taxonomy should be well known, stable, and suitable for correct and consistent
identification by a non-specialist;
2. The biology and general life history should be well understood so as to make ecological
roles known and sources of variation understood;
3. The populations should be readily surveyed and manipulated;
4. Higher taxa (i.e., genera, family) of the indicators should occupy a breadth of habitats
and a broad geographic range so that wide application is possible;
5. At lower taxonomic levels (populations, subspecies, species, etc.), there should be narrow
habitat specialisation so that the ability to detect small geographic differences is provided;
6. The patterns observed in the indicator should actually be an indicator of similar patterns
in other related or unrelated taxa; and
7. A species with potential economic impact may be especially useful for policy purposes
even though it fails to meet other criteria.
While the possibility exists that some indicator species might reliably replace more compre-
hensive species in special cases, the wider application of simple species surrogates seems unlikely.
Taxonomic surrogacy or taxonomic sufficiency (Ellis 1985, Quijón & Snelgrove 2006) is a an
alternative. Taxonomic surrogacy has been effectively treated from a taxonomic perspective by
Bertrand et al. (2006). Irish Sea polychaete data (Mackie et al. 1995) were re-examined at different
taxonomic resolutions employing three equally acceptable phylogenies ranging from splitter influ-
enced to lumper influenced. Good regressions between species richness and family richness existed
for each phylogeny, but slopes were dramatically different. Therefore, the phylogeny used greatly
influences species richness estimates. For most benthic marine fauna, phylogenies are not well
developed.
Field results also suggest caution in the adoption of taxonomic surrogates. Only in the case of
hydrothermal vent fauna have genus, family, and order all been well correlated with species patterns
(Doerries &Van Dover 2003). In deep sediments family-species correlations were poor (Naraya-
naswamy et al. 2003). Quijón & Snelgrove (2006) examined taxonomic surrogacy in a reexamina-
tion of seafloor predator exclusion and found that the family level was effective only when families
contained three or fewer species. They concluded, as with many others, that species-level investi-
gation should be the norm. Following methods used in terrestrial systems (Su et al. 2004), Kar-
akassis et al. (2006) compared similarity analyses of benthic samples in the eastern Mediterranean
with community diversity measured by a broad range of indices. The indicator taxa were multi-
species groups of macrofauna collected by grab, ciliates collected similarly, and megafauna and
fish colleted by trawl. The measures of diversity based on the different indicator groups were poorly
correlated.
Most studies examining taxonomic surrogacy in marine systems have been primarily concerned
with the use of similarity analysis to detect differences rather than estimation of diversity per se.
Warwick (1988) re-examined macrobenthic data from five sites at a coarser resolution, and found
that the family level provided adequate results. Similar sufficiency at the family level has been
found in impacted benthic systems (Olsgard et al. 1998a,b) with the caveat that the level of
resolution should be limited to impacted systems containing steep gradients of impact. Additionally,
family-level studies should only be used following development of a species-level baseline.
The question as to whether one size class can be used to determine diversity trends in another
is especially relevant in benthic ecology due to the traditional separation of macrofauna and
meiofauna studies. Warwick et al. (2006) carried out a carefully designed study across both size
groups with interesting results. Sieve-size fractions of the benthos showed similar diversities when
sampled over a set range of spatial scales. The Shannon Index and Expected Species at a sample
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ROBERT S. CARNEY
size of 50 were used as diversity measures. Diversity of the 63, 125 and 250 µm fractions were
quite similar. Diversities of the 500 and 1000 µm sizes were lower by a factor of about two, but
were similar to one another. No one size fraction could be used as a surrogate for the whole, but
the diversity pattern in the larger and the smaller could possibly be studied at only two sieve sizes.
Taxonomic diversity
The incorporation of taxonomic information into a diversity-like index represents a truly novel
development. Indeed, when the indices that form the taxonomic distinctness approach are examined,
they both stretch and then depart from the traditional view that diversity combines species richness
and proportional abundance. Initially viewed as a need in conservation biology (May 1990, Crozier
1997), the approach has been extensively developed in benthic studies (Clark & Warwick 1998,
1999, 2001, Warwick & Clarke 1998, 2001). Although in use a relatively short time, the approach
is gaining wider acceptance. It has already been reviewed in this journal (Warwick & Clarke 2001),
and is widely available through the PRIMER-5 package of computer analysis routines.
Combination of species diversity measures and numerical taxonomy into a more informative
index was proposed in passing by Sneath & Sokal (1973), but the idea seems to have gone largely
unexplored until conservation biologists sought a means of better assessing diversity (Faith 1992,
Posadas et al. 2001, Mace et al. 2003). In addition to the utility in conservation planning, the concept
is also ecologically appealing as nicely presented by Purvis & Hector (2000). When developing a
operational definition of diversity, three factors rather than two should be included. In addition to
species richness and proportional abundance, we should consider the inherent differences among
the taxa present. Giving a benthic example, we might judge that an assemblage of vermiform
animals consisting solely of polychaetes was in some way less diverse than an assemblage consisting
of burrowing anemones, phoronids, sipunculids, echiurans, holothuroids, and a few polychaetes.
At this time, five descriptors for taxonomic distinctness have been developed (Clark & Warwick
2001; Warwick & Clarke 2001): Taxonomic Diversity ∆, Taxonomic Distinctness ∆*, Average
Taxonomic Distinctness for presence/absence data ∆+, Variation in Taxonomic Distinctness Λ+, and
Total Taxonomic Distinctness s∆+. The first two can be considered three-component diversity indices
combining species richness, proportional abundance, and taxonomic information. The latter three
omit a consideration of abundance. These importance differences are best seen through an exam-
ination of how the measures are calculated.
As introduced in the discussion of Simpson’s λ , the relationship between all pairs of species
can be represented by a symmetrical square matrix (Figure 3). The heart of taxonomic distinctness
is such a matrix of taxonomic distinctness values ωij between each pair. The matrix of distinctness
values is effectively similar to a dendrogram or cladogram. Ideally, ωij values should be based on
carefully developed phylogenies (e.g., Bertrand et al. 2006), but Warwick & Clarke (2001) have
effectively made the case for starting with the Linnaean hierarchy until better values are available.
Unlike phylogenies, the Linnaean hierarchy has fixed ranks. Two individuals in the same species
(i = j) would have a ωij of zero. Two individuals from separate congeneric species (i ≠ j) would
have a ωij of one. If the pair were in confamilial genera, ωij would be two and so on. These
increments can be rescaled to allow for taxonomies with many additional subdivisions such as
tribes, superfamilies, subclasses, etc. (Warwick & Clarke 2001).
The calculation of Taxonomic Diversity and Distinctness combine the values of taxonomic
distinctness with abundance (Equation 4a). For these calculations, each element in the taxonomic
distinctness matrix is weighted by the product of the abundances of each pair of species (xi xj ). The
somewhat more familiar form of ∆ can be made by converting xi xj values to the probability of
encountering the species pair (pij ) simply by dividing each element by N 2 (Equation 4b). The
154
Species × species taxonomic

distinctness weights
Taxonomic hierarchy ωi,j = ωj,i
and distinctness paths
0 ω1,2 ω1,3 ω1,4 ω1,5 ω1,6 … ω1,j
Genus
Family
ω2,1 0 ω2,3 ω2,4 ω2,5 ω2,6 … ω2 ,j
Order
Species 1
ω3,1 ω3,2 ω3,4 ω3,5 ω3,6 … ω3 ,j
Class
0
Phylum
Species 2 ω4,1 ω4,2 ω4,3 0 ω4,5 ω4,6 … ω4 ,j
ω5,1 ω5,2 ω5,3 ω5,4 0 ω5,6 … ω5 ,j
ω6,1 ω6,2 ω6,3 ω6,4 ω6,5 0 … ω6 ,j

Species 3
…
0
ω3,4 = 5
Species 4 ωi,1 ωi,2 ωi,3 ωi,4 ωi,5 ωi,6 … 0
Species 5
Species × species probability of pair

ω6,7 = 3 pi pj = pj pi
Species 6
2
p1 p1 p2 p1 p3 p1 p4 p 1 p5 p1 p6 … p1 pj
p 2 p1
2
p2 p2 p3 p2 p4 p 2 p5 p2 p6 … p2 pj
Species 7
p 3 p1 p3 p2
2
p3 p3 p4 p 3 p5 p3 p6 … p3 pj
Species 8
p 4 p1 p4 p2 p4 p3
2
p4 p 4 p5 p4 p6 … p4 pj
Species 9
p 5 p1 p5 p2 p5 p3 p5 p4 p5
2
p5 p5 … p5 pj
Species 10
p 6 p1 p6 p2 p6 p3 p6 p4 p 6 p5 p62 … p6 pj
…
…
…
Species i
pi p1 pi p2 pi p3 pi p4 pi p5 pi pj … 2
pi
Figure 3 Taxonomic distinctness measures. The taxonomic distinctness suite of indices is based upon deter-
mining distinctness between all pairs of species collected by sampling. As an initial approximation of
phylogenetic relationships, distinctness weight (ω) is half the path length linking a species pair in the taxonomic
hierarchy. The properties of the resulting distinctness matrix can be analyzed and expressed as a purely
taxonomic-distinctness index like ∆*. When combined with a matrix of probabilities of drawing species pairs,
an index of taxonomic diversity (∆) can be obtained that combines species richness, relative abundance and
interspecies evolutionary relationships. This is a major extension of the species diversity concept.
relationship with 1 – λ (Equation 2) explained by Warwick & Clarke (1998) is more obvious in
this presentation. It can also be noted that as N becomes large its effect on the calculated value
quickly becomes small. Seen as an extension of Simpson’s λ , ∆ is the expected or average taxonomic
difference between any pair of specimens drawn from the assemblage on the condition that they
are not the same species.
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ROBERT S. CARNEY
S −1 S
2 ∑∑ω x x
j=2 i< j
ij i j
∆=
( )
Taxonomic Diversity (4a)
N N −1
S −1 S
2 ∑∑ω p p
j=2 i< j
ij i j
∆= (4b)
(1 − N −1 )
S −1 S
∑∑ω x x
j=2 i< j
ij i j
Taxonomic Distinctness ∆* = S S
(5a)
∑∑ x x
j=2 i< j
i j
S −1 S
∑∑ω p p
j=2 i< j
ij i j
∆* = (5b)
1− λ
Taxonomic distinctness, ∆*, is an extension of Taxonomic Diversity based on the ratio of the
product of taxonomic distance and species pair abundances to the same product when all ωij have
been set equal to 1 (Equation 5a). This ratio has the effect of comparing actual weighted taxonomic
distinctness to a reference distance based on all specimens being in the same genus. The relationship
with Simpson’s λ is again more obvious when proportions are used (Equation 5b). Two important
attributes of ∆* are that the ratio eliminates the effects of any scaling that has taken place on the
abundance data, and the direct influence of sample size, n, is eliminated.
When only presence/absence data are available, Taxonomic Diversity and Distinctness reduce
to Average Taxonomic Distinctness. This index is based entirely upon the taxonomic weights and
species richness. Thus, it represents a different definition of diversity than either the index combining
richness with abundance or the three-component definition of Taxonomic Diversity and Distinctness.
S −1 S
+
2 ∑∑ω j=2 i< j
ij
Average Taxonomic Distinctness ∆ = (6)

S ( S − 1)
Excluding studies used in developing the approach, application of the taxonomic distinctiveness
approach is still in the early phases, and much remains to be learned about its utility for answering
a range of questions. Ellingsen et al. (2005) examined its ecological utility by applying the quali-
tative form, ∆+, to soft-sediment macrobenthos at 101 sites along the Norwegian continental shelf
(Ellingsen & Gray 2002). To examine the possibility of surrogacy, annelids, molluscs, and crusta-
ceans were treated separately and then combined for an overall pattern. A distinct gradient of
decreasing values of ∆+ with depth and latitude was found when all taxa were combined, but three
separate groups showed different relationships indicating that no group could serve as a surrogate
156
for the overall pattern. The traditional measure, species richness, showed a modal relationship to
depth and a gradient with latitude and sediment grain size. While ∆+ produced results from the
Norwegian data that warrant additional investigation, these workers concluded that the inconsisten-
cies among taxa may have been an artifact of differences in taxonomic hierarchy rather than ecology.
Aside from ecological applications, conclusions about the applied utility of taxonomic diversity
measures for assessing environmental quality also vary among studies. In the initial application to
a North Sea oil field, the measures showed greater sensitivity than other indices and were monotonic
with the degree of degradation (Warwick & Clarke 1995). Somerfield et al. (1997) found less clear
gradients of impact in a similar system. Applied to three different coastal habitats in Spain and
Portugal (Salas et al. 2006), the measures lacked greater utility than other diversity or habitat-
quality measures. The data from these coastal studies combined grab with hand-collected specimens
and mixed soft bottom, hard bottom, subtidal, and intertidal habitats. There needs to be more
application of the approach across comparable benthic datasets and habitats. This wider application
should be accompanied with refinement by systematists in the method of determining taxonomic
weights before a full critical evaluation is possible.
An unresolved problem with taxonomic distinctness indices is exactly how to interpret them
in the context of diversity theories that largely ignore the phylogenetic aspect of species diversity.
For example, a carefully designed mesocosm experiment examined the combined effects of nutrient
enrichment and physical disturbance on both macrofauna and meiobenthic nematodes (Widdicombe
& Austen 2001, Austen & Widdicombe 2006). Diversity was measured using Total Taxonomic
Distinctness s∆+ (Warwick & Clarke 2001), a measure entirely dependent upon the matrix of
taxonomic distinctness values and species richness. The results for both macrofauna and nematodes
were interpreted as being consistent with the dynamic equilibrium hypothesis of Huston (1979). It
is not clear at this point in the development of taxonomic distinctiveness what Huston’s model
would predict if modified to consider phylogenies. Indeed, it is not obvious how such a modification
should be made other than to accept the unlikely assumptions about generic and familiar competition
and dispersal abilities.
Extrapolation
Benthic surveys typically sample a very small area of bottom and try to characterise the diversity
of a much larger area of sea floor. Collector’s curves of number of species found versus effort
(individuals or samples) seldom approach an asymptote indicating that the complete species inven-
tory has been poorly sampled. Traditionally, benthic ecologists avoided the temptation of extrapo-
lating beyond the actual observed species richness. Conservation biology has, however, driven the
need to extrapolate from samples to much larger areas or to a larger number of samples than actually
taken. Reviews of the methods employed have been written by Bunge & Fitzpatrick (1993) and
Colwell & Coddington (1994) who point out that determining the number of unobserved things is
a statistical challenge in many different fields. The methods are available through the EstimateS
software distributed by Colwell (2005).
Extrapolation in soft-bottom systems has been addressed employing different methods by
Karakassis (1995), Rumohr et al. (2001), Ugland et al. (2003) and Ugland & Gray (2004). Karakassis
employed a method developed from catch statistics identical to earlier work by DeLury (Ugland &
Gray 2003). This method is sample based and plots the observed species at one level of effort, k,
against k + 1. The curve is extrapolated until the two terms are equal. Foggo et al. (2003) applied
the Karakassis method and Rumohr’s modification to beach macrofauna, estuarine oligochaetes
and reef fish. These datasets were modest with the largest for reef fish having 109 samples and
only 33 species. Using the criteria that the predicted species richness should equal the total observed
species richness at 75% sampling effort, it was concluded that different methods gave best estimates
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ROBERT S. CARNEY
at different levels of effort. The Karakassis method always produced low estimates. At high sampling
efforts, the modified Karakassis method gave estimates closer to the actual value.
In addition to Karakassis methods, methods derived from the work of Chao (1984) were
employed in the same comparison. These are non-parametric techniques made popular by the review
of Colwell & Coddington (1994), and are succinctly treated in Magurran (2004). Chao1 and Chao2
are individual-based and sample-based versions of the same relationship (Equations 7a,b). The
estimated species richness is denoted by Ŝ, and the actually observed species richness is Sobs . For
the individual-based case when abundances are known, F1 and F2 denote the number of singleton
and doubleton species. For the sample-based case when only number of occurrences are known;
Q1 and Q2 are the number of species found in just one and two samples.
F2
Chao1 S = Sobs + 1 (7a)
F2
Q2
Chao2 S = Sobs + 1 (7b)
Q2
Concerned that subtidal macrobenthic surveys are typically much more extensive and collect
many more species, Ugland & Gray (2004) carried out a comparison of the Karakassis and Chao
methods using extensive Norwegian shelf data from two regions. One dataset contained 68,298
individuals and 809 species collected in 101 samples (Ellingsen 2001). The second contains more
than three million individuals and 2186 species found in 124 aggregated samples (Ellingsen &
Gray 2002). The first region could be subdivided into five subregions and the second into six based
on various oceanographic factors. Both the Chao and Karakassis methods were found to seriously
underestimate species richness when applied to subsets of data.
Ugland and associates have been pursuing a new application of species accumulation curves
for extrapolation of species richness and bottom heterogeneity. An important advancement was an
independent derivation by Ugland et al. (2003) and Colwell et al. (2004) of an analytical method
of calculating the mean and variance of a sample-based accumulation curve without resorting to
randomisations (Gotelli & Colwell 2001). Ugland et al. (2003) further treated inherent heterogeneity
of the shelf-depth benthos partitioned macrofauna data from Hong Kong and the Norwegian shelf
into subareas and nested the species accumulation curve. Seeking explanations for differences
between the two regions that could be due to bottom heterogeneity, similar nested analyses were
applied to data generated by Arrhenius null models (Ugland et al. 2005) with very good results.
Progress in measurement of diversity over scales: α, β, and γ
One of the most active and interesting areas of diversity research today focuses on the diversity
changes observed when progressing from small to larger spatial scales. These changes are of special
interest because they should reflect the processes through which regional (larger area) dynamics
influence local (smaller area) communities. From a systems perspective, it is a matter of assemblage:
how do smaller units such as communities fit together hierarchically to make a larger unit such as
an ecosystem? When ecologists examine differences in diversity across increasing scales or nested
sets of samples, three general approaches have been taken (Magurran 2004). A value β can be
developed describing the relationship between diversity at one scale α and a larger scale γ.
Diversities can be compared using similarity indices that consider species by species differences.
Finally the species-area relationship can be considered as the area increases.
158
The concept of β diversity was introduced by Whittaker (1960, 1972) who needed a means to
quantify changes in plant diversity along gradients. A scheme was introduced employing ‘inventory’
diversity at four spatial scales and three ‘differentiation’ diversities between adjacent scales. On
the smallest scale is point diversity for a single sampling unit followed by α (habitat), γ (landscape)
and ε (province). The change in diversity between point and α was termed ‘pattern’ diversity. The
most familiar is the difference between α diversity and γ diversity termed β. A simple relationship
between the two is Whittaker’s multiplicative relationship shown in Equation 8 where the denom-
inator is the average α of all components combined to make γ (Magurran 2004).
γ
Whittaker βW = (8)
α
A certain amount of confusion persists surrounding α, β and γ. Most critical confusion is the
distinction between inventory and differentiation diversities. β is not a diversity but is a relationship
between diversities. Since α and γ have the units species, β must be a unitless ratio when the
Whittaker multiplicative approach is used. A related approach is to express β as the slope of species
richness when area sampled or sample number increases (Rosenzweig 1995). In which case β has
the units of species per area or sample number. Wilson & Shumida (1984) evaluated six β diversity
indices, most following the multiplicative tradition, and found the simple Whittaker multiplicative
relationship to most closely meet their criteria, results that were accepted in later reviews (Gray
2000, Magurran 2004).
The second source of confusion is a general inconsistency of terminology in describing scale.
The symbol α is used to describe the diversity of everything from a single sample up to a large
geographic region. In an effort to reduce confusion, Gray (2000) dropped the use of γ and proposed
a nomenclature that built upon a review of benthic system scaling (Thrush et al. 1999). The key
distinction of the system is between the terms habitat and assemblage that should not have restricted
scales versus diversity of points, samples, large areas, and provinces which can be given convenient
set scales. Unfortunately, confusion will be hard to eliminate especially between point and sample
diversity. In oceanographic data archiving the smallest unit, a single core is often recorded as being
a sample rather than the statistical usage in which several cores taken according to a specified
design would comprise a sample. Following the influential evaluation by Lande (1996) of diversity
measures, Crist & Veech (2006) proposed simply using αi to denote diversity at all levels with the
actual level specified by the subscript. Further each level αi is composed of a set of lower level
values. Effectively, no fixed scales are used, and it is the burden of the investigator to specify the
scales over which αi values are nested.
As part of the renewed interest in the scales of diversity and how large ecosystems fit together,
it has become appreciated that there is no one right way of envisioning β diversity. A most important
recent development is the use of additive rather than multiplicative measures, i.e., γ is the sum of
α and β rather than their product. The general relationship attributed to Lande (1996) is shown in
Equation 9. Development of an actual computational form is more complicated. Veech et al. (2002)
traced the origins of additive diversity partitioning to MacArthur (1966) and Levins (1968), noting
that these initial works did not apply the same α, β and γ symbols and terminology as Whittaker
(1960), whereas Lande (1996) did. Adopting additive partitioning, β diversity can be seen as the
difference between the average diversity of sub units and the overall diversity of the set they are
in (Loreau 2000).
Additive Diversity γ = α+β (9)
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ROBERT S. CARNEY
The first application of additive diversity partitioning to the benthos was not actually presented
as such. Ugland et al. (2003) developed an approach to accumulation curve analysis — the analytical
species accumulation (ASA) approach, which is based upon accumulation within subsets of samples
and a novel analytical expression for the accumulation curve rather than Monte Carlo simulations.
The method was used to extrapolate total species richness in areas of the Norwegian shelf and
Hong Kong harbour. Crist & Veech (2006) recognised, however, the additive nature of the rela-
tionship in ASA between observed species richness in a combined set and the average species
richness of the members of the subset. On further investigation of the additive and multiplicative
models of β diversity, Kiflawi & Spencer (2004) established the interrelationship βM = βA/α and
explored the statistical properties of both approaches. No evidence of the benefit of one over the
other was presented.
The analysis of similarity to investigate β and for other purposes is so extensively used in
benthic studies (Gray 2000) that it is beyond the scope of this review. The partitioning of a similarity
matrix to study β diversity has been compared to the use of the variance of a raw species by a
sample array (Legendre et al. 2005).
As previously noted a goal of assessing diversity changes across scales and across system
hierarchies is to gain information about processes. A seminal work of this kind was based on a
study of West Indian bird communities (Terborgh & Faaborg 1980). A simple relationship was
proposed that distinguished between species saturated and unsaturated communities. Saturated
communities were those having such strong species interactions that no new species from the
regional pool could successfully enter. Unsaturated communities lacked similarly intense interaction
allowing additional species to enter from the larger pool. The difference between saturated and
unsaturated communities should be detected by simple plots from many localities and regions of
local (α) versus regional species richness (γ). Saturated communities would show an asymptote
while unsaturated would show a linear relationship. Unfortunately, this simple scheme has failed
in a large number of terrestrial and marine studies (Russell et al. 2006). The determination of local
versus regional diversity is, however, seen as an important task in diversity studies assuming
adequate attention is paid to designs that make appropriate comparisons (Ricklefs 2004).
Ellingsen & Gray (2002) carried out an examination of diversity at different scales that
employed four approaches to β and examined the relationship between local (α) and regional (γ)
diversities. The smallest scale was represented by five pooled van Veen grabs taken at 101 sites
along the length of the Norwegian continental shelf. Whittaker’s βw, number of species shared
between all pairs, biotic distinctness and Bray-Curtis similarity were calculated. Regional pooling
of data produced γ diversities. While the results were discussed in the context of lacking a latitudinal
gradient, other points were equally important. β values were found to vary with taxa such that no
group could serve as a surrogate for the whole. α was found to bear no clear relationship with γ
that would indicate strong regional control of local diversity. The Bray-Curtis values and biotic
distinctness, both similarity measures, reflected γ diversity changes more than βw .
Biogeographic studies that compare diversity across large scales can differ in the manner in
which diversity at the largest scale (γ) is determined. It can be arrived at by pooling data into larger
and larger composites. When this is done there may be relationships between α and λ diversities
that reflect the pooling process rather than ecology. Therefore an independent estimate of large-
scale diversity is desirable. To accomplish this a list of regional species can be compiled independent
of the smaller-scale sampling by using published lists, biogeographic archives, museum collections,
etc. In the case of well-studied areas such as the Norwegian shelf the regional pool of macrofaunal
species should be especially well known. Unfortunately, few similarly well-known regions exist.
Developing regional pools from multiple sources of taxonomic knowledge also has problems
associated with it. Such regional species lists may fail to draw important distinctions between
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habitats in heterogenous regions, overestimating the number of species available for colonisation
and survival at small-scale sites (Russell et al. 2006). Traditional information may be expressed
only in terms of depth and geographic ranges. The limits of observed occurrence set the boundaries,
and it is implicit that the species is in the regional pool throughout the range when discontinuous
distribution is more often the actual case (Hurlbert & White 2005). Compiled ranges when strong
boundaries exist such as the surface and the maximum depth of the ocean can be expected to
produce maxima in diversity that may have no ecological relevance, the Mid-Domain effect (Colwell
et al. 2005, Connolly 2005). Pineda & Caswell (1998) examined species richness and ranges on
the Gay Head-Bermuda data and found elements of agreement and disagreement between model
and observed diversity.
Examining large-scale patterns

Study of large-scale patterns is an extremely active area of benthic diversity research (Gray 2002).
The global latitude gradient is the primary focus due to the large number of taxonomists and
systematists addressing that question. A smaller number of experts continue to study species
distribution and diversity on the continental margins over the transition from shallow to abyssal
depths. These gradients are being examined from both a local and a regional perspective. The much
shorter spatial scale of the bathymetric gradient makes small-scale analysis easier with diversity
being first measured sample-by-sample. The great expanse of latitude makes large-scale range
compilations taxa-by-taxa based on sampling and archived records the most common approach.
Simple species richness is the most often used measure of diversity.
Global latitude gradient

There has been important progress in synthesising the accumulated knowledge about distributions
in the ocean. In a meta-analysis of 232 published studies including 102 of coastal benthos and 34
of the deep-sea, it was established that marine species richness increases towards the equator
(Hillebrand 2004a,b). Marine diversity shows this latitude effect as strongly as terrestrial. Regional
(γ) diversity shows the effect more strongly than local (α). Furthermore, the actual measure of
diversity used did not influence the correlation between diversity and latitude. It did, however,
influence the slope of the gradient, being steepest when species richness was used. Within this
global pattern, particular habitats and faunal groups did vary. Among the faunal categories consid-
ered, epibenthic and endobenthic gradients were significant but weaker than others. Within marine
habitats, the correlation between diversity and latitude is stronger for the deep-sea than coastal
waters. The slope of the gradient was steeper for coastal habitats, but the shallow/deep difference
was not statistically significant.
These generalities about the marine latitude gradient may prove valid with additional investi-
gation, but should be considered with caution. They were generated from such a large and diverse
literature by means of meta-analysis, a procedure for combining results from multiple studies widely
used in medicine, education, and social research (Hedges & Olkin 1985), that is being applied to
ecological research (Gurevitch & Hedges 1993, Rosenberg et al. 2000). Meta-analysis can be a
powerful tool when seeking consensus from an extensive body of literature with contradictory
findings. Unfortunately, in meta-analysis the individual studies become anonymous along with all
their assumptions, incompatibilities, and possible errors (Slavin 1986). Given the weakness of the
latitudinal effect for epi- and endobenthos, more information about the contributing studies would
be informative. The actual latitudinal patterns of the soft bottom still seem poorly studied. So much
so that the observation by Clarke & Crame (1997) of a lack of convincing evidence of a soft bottom
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ROBERT S. CARNEY
(α) diversity cline may still be valid. As noted by Hillebrand, many individual studies did not find
significant gradients.
A persistent problem with the assessment of global diversity patterns is that the spatial scale
of the underlying data is often unknown and most likely mixed. This was a problem even in the
earliest conflicting findings. When Thorson (1957) first proposed the pattern of a benthic epifauna
diversity maximum in the tropics contrasted by a more geographically uniform infauna diversity,
he was speaking mainly in terms of regional (γ) diversity compiled from large studies. When Sanders
(1968) countered that infauna also show highest diversities on the tropical shelf, his supporting
data were based on sample (α) diversity.
Global comparisons of sample (α) diversity based on compilations of multiple studies are quite
difficult due to lack of habitat comparability, non-standard methods, and inconsistent taxonomy.
All of these factors may bias meta-analysis results. A narrower and more refined analysis was
undertaken by Attrill et al. (2001) making appropriate comparisons by a careful screening of studies
of intertidal mudflats. Twenty such studies were selected based upon a restricted salinity and median
grain size range, comparable samplers, and use of a 500-µm sieve. Diversity was measured with
Simpson’s index based on performance criteria suggested by Rozenweig (1995). Fisher’s (α) was
rejected due to a lack of fit of the log-series for many of the datasets. Based on this high-quality
data, an increase in diversity was found from high latitudes towards the equator. A similar approach
could be applied to other habitats if datasets become available.
Ideally, global gradients would best be determined through co-ordinated global sampling
designed to examine a range of scales. An example of what can be accomplished is seen in the
survey of subtidal rock wall epibiota by Witman et al. (2004). Smallest-scale species richness,
extrapolated richness, and Choa2 estimates of full species richness were determined from photo
transects on subtidal rock walls at twelve global sites. This meets the criteria that similar habitats
be studied and scales be specified. Regional diversity was independently estimated on the basis of
local species lists and experts. Both local and regional diversity increased toward the equator with
higher latitudes having a greater per cent of the regional pool found in local samples. A more
modest approach that controlled the uniformity of habitat to some degree was taken by Gobin &
Warwick (2006) by putting artificial substrate at four regions from 10°N to 63°S. Shannon’s index
and others were measured. Place to place differences were found, but neither polychaetes nor
nematodes conformed to a latitudinal gradient. Similar global sampling could be conducted for
infauna.
Many large-scale studies are taxonomically restricted. While these studies seldom claim that
the targeted taxon serves as a surrogate for total community diversity, surrogacy is often implied
in the discussion of the theoretical consequences of the results. One of the best-examined compo-
nents of global-scale diversity are the molluscs (Rex et al. 2004), and have the added benefit of
speciation and extinction rates estimated from the fossil record (Jablonski et al. 2006). Many taxa-
restricted studies omit fine-scale assessment and determine larger-species richness from range
compilations. A good example is the study by Roy et al. (2000) of 930 marine bivalves distributed
on the eastern Pacific continental shelf between 71°N and 5°S. A strong latitudinal gradient with
maximum species richness at about 10°N latitude was found. There was a good correlation between
diversity and surface temperature. Taxa-restricted studies are often in conflict when different habitats
and regions are studied. When Valdovinos et al. (2003) extended the molluscs study southward, a
poleward increase in species richness was found but no correlation with surface temperature.
Similarly, the infaunal protobranchs that showed no latitude gradient on the northeast Pacific shelf
were shown to have such a gradient in the deep Atlantic (Allen & Sanders 1996, Rex et al. 1997,
2004).
162
Deep-sea diversity
Studies of large-scale diversity patterns in the deep sea continue to be hampered by a relative
paucity of samples and a large backlog of undescribed species. Sampling is increasing, however,
with the advent of deep-sea resource development, and a core of taxonomists who continue to make
progress in species description. There are actually three benefits of the paucity of samples. First,
diversity is usually measured on a fine-scale sample-by-sample basis, and then compiled for larger-
scale analysis. Second, so few sampling devices have been used that scales are well known. Third,
the same experts have often been able to study many separate studies, assuring a high level of
taxonomic comparability. The main large-scale gradients of interest are latitude and depth. There
is also an interest in estimating the total species richness of this vast region. A variety of diversity
tools have been used. Species richness is usually rarefied, and indices such as Simpson’s and
Shannon’s are often used.
A latitude gradient with increased tropical diversity has been reported for isopods (Poore &
Wilson 1993), Foraminifera (Culver & Buzas 2000), polychaetes (Paterson & Lambshead 1995),
bivalve molluscs (Rex et al. 2000) and cumaceans (Gage et al. 2004). Local and regionally compiled
species richness combined with a regression was used to reach these conclusions. The analysis of
Culver & Buzas (2000) was distinct and used species count data from more than 110 samples in
published studies. Species richness, and Fisher’s α were calculated. Clear gradients were significant
in a regression of these parameters against latitude for both measures of diversity. Wilson (1998)
carried out isopod work restricted to the Atlantic, using Expected Species with rarefaction to 200
individuals on 66 samples. The isopods were partitioned into the Flabellifera and Asellota. The
former showed negative correlation with depth and latitude. The latter showed positive correlation
with only depth. Evolutionary history was thought to still exert a strong control over broad-scale
patterns.
The idea that the deep sea is species rich with a maximum diversity at some middle depth on
the continental margin originated with Sander’s (1968) observation and is now reasonably well
demonstrated around the north Atlantic (Rex et al. 1997). The generality of the pattern in the global
ocean and across taxa is still open to valid questions (Gray 2001). In effect, there will have to be
many better-designed shelf to abyss sampling programmes around the world to settle the matter.
The mid-slope modal pattern can be viewed as unexpected from two perspectives. First,
population sizes decrease progressively with depth due to loss of nutrient value of detritus as it
sinks from its photosynthetic origins. Second the deep-sea bottom appears to become progressively
more homogenous and possibly niche poorer with depth. The subject has received extensive recent
review (Gray 2001, Levin et al. 2001, Snelgrove & Smith 2002, Tyler 2003) and a comprehensive
book is in progress (Rex personal communication). Most analyses have been based on rarefacted
species richness of samples and focus questions on diversity maintenance on small scales. Rex
et al. (2005) examined compiled mollusc depth ranges assessing both species richness within depth
bands and noting the range widths and end points. This approach to diversity analysis lead to the
hypothesis that the abyssal region is a sink largely populated by species with larger populations at
shallower depths on the slope.
Deep-sea diversity has also been controversial with respect to global extrapolation of total
marine species richness. Extrapolating the species accumulation curve generated from samples
collected on the Atlantic continental slope of the United States, Grassle & Maciolek (1992) predicted
a global marine species richness of the order of 108 species with most residing in deep water. This
hyper diversity was quickly challenged on grounds of methodology (May 1992), and in light of
contrary benthic diversity data (Gage & May 1993, Poore & Wilson 1993, Gray 1994). The notion
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ROBERT S. CARNEY
has persisted, however, that the deep sea may be exceptionally diverse especially if poorly resolved
groups like the nematodes were better studied.
The issue of deep hyperdiversity has been specifically addressed and rejected in the case of
nematodes by Lambshead & Boucher (2003) employing extrapolation methods. As a caution against
extrapolating far beyond a region of sampling, it was shown that the extrapolation based on north-
south accumulation gave different results than accumulation East-West. Having established the
tentative nature of the conclusions, accumulation curves and Chao methods were used to estimate
total nematode diversity in 16 ocean regions over a wide depth range. The deep regions did not
have an exceptionally high estimated diversity of nematode fauna. The question then revolves
around the issue of local versus regional diversity and whether the deep sea substantially differed
from shallower depths. At depth α diversity may be high while γ is comparably low due to a
widespread but very patchy species pool across the great expanse of the abyssal plain.
Conclusions
In undertaking the historical review I expected that early benthic ecologists would have expressed
an interest in the variety of animals only to be hindered by a lack of analytical tools. Except for
the mention of higher variety in deep water CHALLENGER samples (Murray 1895) and a criticism of
the lack of relative abundance data (Stuxburg 1883), there was limited interest. Subsequent adoption
of the rigid early community concept reduced the interest even more. As wrongly noted by Allee
(1934), characterisation of the representative species was such a sufficent approach that details
about the minor constituents were unnecessary; in effect total diversity was seen as irrelevant!
There are, however, at least two lessons to be learned from history about species distributions and
the parsimonious description of a community or assemblage. Thorson’s (1957) criteria for the
selection of characteristic species and the four types of characteristic species that could be consid-
ered. First, the nature of the four types (first order, second order, third order, and influent) makes
it clear that early workers recognised the relationship between geographic range and occurance in
samples. The first-order species in a community were common but geographically restricted. The
influents (fourth order) were common but found over too large a range to be useful in characterising
a specific community found within that range. This is quite similar to the suggestion by Ugland et al.
(2005) that benthic assemblages might be best considered a combination of a widely and narrowly
distributed species. In future investigations small-scale diversity should be partitioned according
to the larger ranges of the contributing species. Second, is the matter of parsimony. Historically
characteristic species were used to name and map assemblages. Scientifically, there may be little
value in the naming of assemblages. Much of the demand for biodiversity information, however,
falls into the area of policy and political science. In this arena, it may be best to return to a system
of assemblage nomenclature that first describes and maps assemblages in terms of characteristic
species. Then, when greater scientific characterisation is needed, composition and diversity can be
appropriately quantified. Certainly some improvement over “Maldane sarsi-Ophiura sarsi commu-
nity” (Thorson 1957) is now possible, but equally simple and descriptive nomenclature should be
of great utility when conservation is debated.
Benthic ecologists face the issue of which diversity measure to use. Lande’s (1996) criteria that
good measures of diversity should be non-parametric is widely accepted throughout ecology. Judging
by the most common choice of methods, most benthic ecologists prefer to use non-parametric
indices rather than fit a distribution to species abundance data. Nevertheless, the log-series has
proven effective in application to benthic foraminiferans and is strongly advocated as a null model
(Buzas & Hayek 2005). Similarly, the log-normal has proven effective in the study of macrobenthos
and has been effectively advocated as a null model (Gray et al. 2006a,b). The great advantage of
using a parametric distribution is that the full complexity of the data may be most parsimoniously
164
described without the ambiguity of an index that combines two or more aspects of the sample.
Therefore, it is valuable to retain the use of distributions especially when previous work on the
same faunal components and in the same region have shown there to be a good and consistent fit.
Non-parametric indices should be the preferable means of describing diversity when the data
considered are from previously unstudied areas or faunal components. Similarly, when the spatial
and temporal scale of sampling are so great as to include more than a single assemblage type, non-
parametric indices are preferred.
In his influential book Huston (1994) gave a brief but adequate discussion of diversity measures
and offered the suggestion that a lot of exploration of these tools produced more mathematics than
understanding of actual biological diversity. This may have seemed the case at the time, but some
developments with origins in benthic ecology are truly innovative. Taxonomic distinctness and the
associated measures of phylogenetic variability are the primary example. When the measure com-
bines phylogenetic information with species richness and abundance as in the case of taxonomic
diversity, it becomes an extension of how biological diversity is conceptualised. Once abundance
is removed, however, it is probably best that these measures should not be treated as diversity in
order to minimise confusion. The study of taxonomic distinctness is an endeavour in itself, and
determining patterns across many marine habitats and taxa is an exciting new undertaking. Similarly,
extrapolation from a small area of sea floor actually sampled to areas being considered for conser-
vation management is an important advancement.
The impact of computer technology on the study of diversity patterns cannot be overstated.
However, this calls into question why ecologists still employ diversity analyses developed prior to
easy access to digital computers? Indices like Simpson’s and Shannon’s are products of the
mechanical calculator age when characterising fauna samples with more than a single value was a
lengthy process. When patterns are found, they have to be dissected to be understood. How did
species richness and relative abundance change? Scientifically, multiple measures should be used
beginning with species richness, rarefacted values to allow for better comparisons, and then indices
giving different weights to different fractions. In effect, ease of computation has already made this
the standard approach in benthic ecology. In the policy area, however, only one or very few measures
should be used. Species richness with an adjustment for sample size is the easiest to explain to a
non-scientific audience.
While the term informatics may seem like unnecessary jargon, having access to vast amounts
of information through networked data sources is an extremely powerful tool. This is seen in
Hillebrand’s succinct summation of latitudinal trend in the ocean and on land through the use of
literature keyword searches and meta-analysis. A traditional scholarly review could only have
produced a list of contradictory findings. There is, however, a limit to the utility of reviews,
traditional or otherwise. Access to real data is preferable. Currently, there are several international
efforts to create open data archives. Many databases are best suited for species-by-species investi-
gation of occurrences, and will require additional development to be useful in diversity analyses
that begin core by core. Once suitable formats are available, it can be hoped that complete regional
surveys will be made available for re-examination.
The importance of the assessment of diversity across multiple scales will continue to increase.
Conservation policy needs information on the location of diversity hot spots and the spatial extent
of relatively homogenous assemblages. Theoretical and applied ecology needs information on how
regional species pools influence assemblages on local scales. Development of new cross-scale
diversity measures is an active and needed area of research. Unfortunately, ambiguity as to what
scales α and γ precisely refer to has made published results hard to interpret. Adhering to the niche
theory view that local diversity must be influenced by competition and partitioning of resources,
species count data should be collected, analyzed and initially reported on a consistently small scale.
Benthic ecologists have a great advantage in this regard as grabs and corers are limited to less than
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0.5 m2. These point measurements can then be pooled into the larger sets allowed by sampling
design to cover larger scales. Rather than depend too much on nomenclature, the scales should
always be specified.
As a final observation, there must be more large-scale studies conducted at locations carefully
selected to test the effects of ocean processes upon diversity across a wide range of scales. Standard
methods must be used and comparable taxonomy assured. Such well-designed studies will provide
for a better understanding than re-examination of old archived data far beyond the level of inquiry
anticipated in the original sampling designs. Ideally, scientific questions about ocean diversity are
adequate to drive such ambitious new sampling and analysis. More realistically, monetary support
is more likely to be mandated by resource mangers in the policy arena if diversity studies can be
shown to be an indispensable management tool providing readily understood results.
References
Allee, W.C. 1923. Studies in marine ecology. I. The distribution of common littoral invertebrates of the Woods
Hole region. Biological Bulletin 36, 96–104.
Allee, W.C. 1934. Concerning the organization of marine communities. Ecological Monographs 4, 541–554.
Allen, J.A. & Sanders, H.L. 1996. The zoogeography, diversity and origin of the deep-sea protobranch bivalves
of the Atlantic: the epilog. Progress in Oceanography 38, 95–153.
Andelman, S.J. & Fagan, W.F. 2000. Umbrellas and flagships: efficient conservation surrogates or expensive
mistakes? Proceedings of the National Academy of Sciences U.S.A. 97, 5954–5959.
Anderson, A.N. 1995. Measuring more of biodiversity: genus richness as a surrogate for species richness in
Australian ant faunas. Biological Conservation 73, 39–43.
Attrill, M.J., Stafford, R. & Rowden, A. 2001. Latitudinal diversity patterns in estuarine tidal flats: indications
of a global cline. Ecography 24, 318–324.
Austen, M.C. & Widdicombe, S. 2006. Comparison of the response of meio- and macrobenthos to disturbance
and organic enrichment. Journal of Experimental Marine Biology and Ecology 330, 96–104.
Bertrand, Y., Pleijel, F. & Rouse, G.W. 2006. Taxonomic surrogacy in biodiversity assessments, and the
meaning of Linnean ranks. Systematics and Biodiversity 4, 149–159.
Bunge, J. & Fitzpatrick, M. 1993. Estimating the number of species: a review. Journal of the American
Statistical Association 88, 364–372.
Buzas, M.A. & Culver, S.J. 1989. Biogeographic and evolutionary patterns of continental-margin benthic
foraminifera. Paleobiology 15, 11–19.
Buzas, M.A. & Hayek, L.C. 2005. On richness and evenness within and between communities. Paleobiology
31, 199–220.
Carney, R.S. 2005. Zonation of deep-sea biota on continental margins. Oceanography and Marine Biology:
An Annual Review 43, 211–278.
Chao, A. 1984. Non-parametric estimation of the number of classes in a population. Scandinavian Journal of
Statistics 11, 265–270.
Clarke, A. & Crame, J.A. 1997. Diversity, latitude and time: patterns in the shallow sea. In Marine Biodiversity:
Patterns and Processes, R.F.G. Ormond et al. (eds). Cambridge: Cambridge University Press, 122–147.
Clarke, K.R. & Warwick, R.M. 1998. A taxonomic distinctness index and its statistical properties. Journal of
Applied Ecology 35, 523–531.
Clarke, K.R. & Warwick, R.M. 1999. The taxonomic distinctness measure of biodiversity: weighting the step
lengths between hierarchical level. Marine Ecology Progress Series 184, 21–29.
Clarke, K.R. & Warwick, R.M. 2001. A further biodiversity index applicable to species lists: variation in
taxonomic distinctness. Marine Ecology Progress Series 216, 265–278.
Clarke, R.B. & Milne, A. 1955. The sublittoral fauna of two sandy bays on the isle of Cumbrae, Firth of
Clyde. Journal of the Marine Biological Association of the United Kingdom 34, 161–180.
Clements, F.E. & Shelford, V.E. 1939. Bio-ecology. New York: John Wiley and Sons.
166
Colwell, R.K. 2005. EstimateS Users Guide. Online. Available HTTP: http://purl.oclc.org/estimates (accessed
31 October 2006).
Colwell, R.K. & Coddington, J.A. 1994. Estimating the terrestrial biodiversity through extrapolation. Philo-
sophical Transactions of the Royal Society of London Series B 345, 101–118.
Colwell, R.K., Mao, C.X. & Chang, J. 2004. Interpolating, extrapolating and comparing incidence-based
species accumulation curves. Ecology 85, 2717–2727.
Colwell, R.K., Rahbek, C. & Gotelli, N.J. 2005. The mid-domain effect: there’s a baby in the bathwater.
American Naturalist 166, 149–154.
Connolly, S.R. 2005. Process-based models of species distributions and the mid-domain effect. American
Naturalist 166, 1–11.
Costello, M.J. & Berghe, E.V. 2006. ‘Ocean biodiversity informatics’: a new era in marine biology research
and management. Marine Ecology Progress Series 316, 203–214.
Costello, M.J., Bouchet, P., Emblow, C.S. & Legakis, A. 2006. European marine biodiversity inventory and
taxonomic resources: state of the art and gaps in knowledge. Marine Ecology Progress Series 316,
257–268.
Crist, T.O. & Veech, J.A. 2006. Additive partitioning of rarefaction curves and species-area relationships:
unifying α-, β-, and γ-diversity with sample size and habitat area. Ecology Letters 9, 923–932.
Crozier, R.H. 1997. Preserving the information content of species: genetic diversity, phylogeny, and conser-
vation worth. Annual Review of Ecology and Systematics 28, 243–268.
Culver, S.J. & Buzas, M.A. 2000. Global latitudinal species diversity gradient in deep-sea benthic Foraminifera.
Deep-Sea Research I 47, 259–275.
Doerries, M.B. & Van Dover, C.L. 2003. Higher-taxon richness as a surrogate for species richness in chemo-
synthetic communities. Deep-Sea Research I 50, 749–755.
Eduardo, C. & Grelle, V. 2002. Is higher-taxon analysis a useful surrogate for species richness in studies of
neotropical mammal diversity? Biological Conservation 108, 101–106.
Ellingsen, K.E. 2001. Biodiversity of a continental shelf soft-sediment macrobenthic community. Marine
Ellingsen, K.E., Clarke, K.R., Somerfield, P.J. & Warwick, R.M. 2005. Taxonomic distinctness as a measure
of diversity applied over a large scale: the benthos of the Norwegian continental shelf. Journal of
Animal Ecology 74, 1069–1079.
Ellingsen, K.E. & Gray, J.S. 2002. Spatial patterns of benthic diversity: is there a latitudinal gradient on the
Norwegian continental shelf? Journal of Animal Ecology 71, 373–389.
Ellis, D. 1985. Taxonomic sufficiency in pollution assessment. Marine Pollution Bulletin 16, 459–459.
Fabri, M.C., Galeron, J., Larour, M. & Maudire, G. 2006. Combining the Biocean database for deep-sea
benthic data with the online Ocean Biogeographic Information System. Marine Ecology Progress
Series 316, 215–224.
Faith, D.P. 1992. Conservation evaluation and phylogenetic diversity. Biological Conservation 61, 1–10.
Fisher, A.A., Corbert, A.S. & Williams, C.B. 1943. The relationship between the number of species and the
number of individuals in a random sample of an animal population. Journal of Animal Ecology 12, 42–58.
Foggo, A., Attrill, M.J., Frost, M.T. & Rowden, A.A. 2003. Estimating marine species richness: an evaluation
of six extrapolative techniques. Marine Ecology Progress Series 248, 15–26.
Forbes, E. 1859. Map of the distribution of marine life, illustrated chiefly by fishes, molluscs and radiata
showing also the extended limits of the homoiozoic belts. In The Physical Atlas of Natural Phenomena,
A.K. Johnson (ed.). Edinburgh: W. & A.K. Johnson.
Ford, E. 1923. Animal communities of the level sea bottom in the waters adjacent to Plymouth. Journal of
the Marine Biological Association of the United Kingdom 13, 165–224.
Gage, J. 1972. Community structure of benthos in Scottish sea-lochs. 1. Introduction and species diversity.
Gage, J., Lambshead, P.J.D., Bishop, J.D.D., Stuart, C.T. & Jones, N.S. 2004. Large-scale biodiversity patterns
of Cumacea (Peracarida: Crustacea) in the deep Atlantic. Marine Ecology Progress Series 277, 181–196.
Gage, J. & May, R.M. 1993 Biodiversity — a dip into the deep-seas. Nature 365, 609–610.
Gage, J. & Tett, P.B. 1973. Use of log-normal statistics to describe benthos of lochs Etive and Creran. Journal
of Animal Ecology 42, 373–382.
167
ROBERT S. CARNEY
Gaston, K.J. & Williams, P.H. 1993. Mapping the world’s species — the higher taxon approach. Biodiversity
Letters 1, 2–8.
Gobin, J.F. & Warwick, R.M. 2006. Geographical variation in species diversity: a comparison of marine
polychaetes and nematodes. Journal of Experimental Marine Biology and Ecology 330, 234–244.
Gorelick, R. 2006. Combining richness and abundance into a single diversity index using matrix analogues
of Shannon’s and Simpson’s indices. Ecography 29, 525–530.
Gotelli, N.J. & Colwell, R.K. 2001. Quantifying biodiversity: procedures and pitfalls in the measurement and
comparison of species richness. Ecological Letters 4, 379–391.
Grassle, J.F. & Maciolek, N.J. 1992. Deep-sea species richness: regional and local diversity estimates from
quantitative bottom samples. American Naturalist 139, 313–341.
Gray, J.S. 1981a. The Ecology of Marine Sediments: An Introduction to the Structure and Function of Benthic
Communities. Cambridge: Cambridge University Press.
Gray, J.S. 1981b. Detecting pollution induced changes in communities using the log-normal distribution of
individuals among species. Marine Pollution Bulletin 12, 173–176.
Gray, J.S. 1983. Use and misuse of the log-normal plotting method for detection of effects of pollution — a
reply. Marine Ecology Progress Series 11, 203–204.
Gray, J.S. 1985. Ecological theory and marine pollution monitoring. Marine Pollution Bulletin 16, 224–227.
Gray, J.S. 1994. Is deep-sea species-diversity really so high: species-diversity of the Norwegian continental
shelf. Marine Ecology Progress Series 112, 205–209.
Gray, J.S. 2000. The measurement of marine species diversity, with an application to the benthic fauna of the
Norwegian continental shelf. Journal of Experimental Marine Biology and Ecology 250, 23–49.
Gray, J.S. 2001. Marine diversity: the paradigms in patterns of species richness examined. Scientia Marina
65 (Suppl. 2), 41–56.
Gray, J.S. 2002. Species richness in marine sediments. Marine Ecology Progress Series 244, 285–297.
Gray, J.S., Bjørgesæter, A., Ugland, K.I. & Frank, K. 2006a. Are there differences in structure between marine
and terrestrial assemblages. Journal of Experimental Marine Biology and Ecology 330, 19–26.
Gray, J.S., Bjørgesæter, A., Ugland, K.I. & Frank, K. 2006b. On plotting species abundance distributions.
Journal of Animal Ecology 75, 752–756.
Gurevitch, J. & Hedges, L.V. 1993. Meta-analysis: combining the results of independent experiments. In
Design and Analysis of Ecological Experiments, S.M. Scheiner & J. Gurevitch (eds). New York:
Chapman Hall, 378–398.
Hamilton, A.J. 2005. Species diversity or biodiversity? Journal of Environmental Management 75, 89–92.
Harper, J.L. & Hawskworth, D.L. 1994. Biodiversity: measurement and estimation, preface. Philosophical
Transactions of the Royal Society London Series B 345, 5–12.
Hayek, L.C. & Buzas, M.A. 1997. Surveying Natural Populations. New York: Columbia University Press.
Hedgepeth, J.W. 1957. Concepts of marine ecology. In Treatise on Marine Ecology and Paleoecology, J.W.
Hedgepeth (ed.). The Geological Society of America Memoir 67, 29–52.
Hedges, L.V. & Olkin, I. 1985. Statistical Methods for Meta-Analysis: Orlando, Florida: Academic Press.
Hill, M. 1973. Diversity and evenness: a unifying notation and its consequences. Ecology 54, 427–432.
Hillebrand, H. 2004a. Strength. Slope and variability of marine latitudinal gradients. Marine Ecology Progress
Series 273, 251–267.
Hillebrand, H. 2004b. On the generality of the latitudinal diversity gradient. American Naturalist 163, 192–211.
Holme, N.A. 1966. Bottom fauna of the English Channel. 2. Journal of the Marine Biological Association of
the United Kingdom 46, 401–493.
Hubbell, S.P. 2001. The Unified Neutral Theory of Biodiversity and Biogeography. Princeton, New Jersey:
Princeton University Press.
Hurlbert, A.H. & White, E.P. 2005. Disparity between range map- and survey-based analyses of species
richness: patterns, processes and implications. Ecology Letters 8, 319–327.
Hurlbert, S. 1971. The non-concept of species diversity: a critique and alternative parameters. Ecology 52,
577–586.
Huston, M.A. 1979. A general hypothesis of species diversity. American Naturalist 113, 81–101.
Huston, M.A. 1994. Biological Diversity: The Coexistence of Species on Changing Landscapes. Cambridge:
Cambridge University Press.
168
Jablonski, D., Roy, K. & Valentine, J.W. 2006. Out of the tropics: evolutionary dynamics of the latitudinal
diversity gradient. Science 314, 102–106.
Jones, N.S. 1950. Marine bottom communities. Biological Reviews 25, 283–313.
Karakassis, I., Machias, A., Pitta, P., Papadopoulou, K.N., Smith, C.J., Apostolaki, E.T., Giannoulaki, M.,
Kouysoubas, D. & Somarakis, S. 2006. Cross-community congruence of patterns in a marine ecosys-
tem: do the parts reflect the whole? Marine Ecology Progress Series 310, 47–54.
Karakassis, I. 1995. Sinfinity: a new method for calculating macrobenthic species richness. Marine Ecology
Kiflawi, M. & Spencer, M. 2004. Confidence intervals and hypothesis testing for beta diversity. Ecology 85,
285–2900.
Lambshead, P.J.D. & Boucher, G. 2003. Marine nematode deep-sea biodiversity — hyperdiverse or hype?
Journal of Biogeography 30, 475–485.
Lande, R. 1996. Statistics and partitioning of species diversity, and similarity among multiple communities.
Oikos 76, 5–13.
Lande, R., DeVried, P.J. & Walla, T.R. 2000. When species accumulation curves intersect: implications for
ranking diversity using small samples. Oikos 89, 601–605.
Laptikhovsky, V. 2006. Latitudinal and bathymetric trends in egg size variation: a new look at Thorson’s and
Rass’s rules. Marine Ecology 27, 7–14.
Legendre, P., Borcard, D. & Pere-Neto, P.R. 2005. Analyzing beta diversity: partitioning the spatial variation
of community composition matrix. Ecological Monographs 75, 435–450.
Levin, L.A., Etter, R.J., Rex, M.A., Gooday, A.J., Smith, C.R., Pineda, J., Stuart, C.T., Hessler, R.R & Pawson,
D. 2001. Environmental influences on regional deep-sea species diversity. Annual Review of Ecology
and Systematics 132, 51–93.
Levins, R. 1968. Evolution in Changing Environments. Princeton, New Jersey: Princeton University Press.
Lie, U. 1974. Distribution and structure of benthic assemblages in Puget Sound, Washington, U.S.A. Marine
Biology 26, 203–223.
Lie, U. & Kelly, J.C. 1970. Benthic infauna communities off coast of Washington and in Puget Sound —
identification and distribution of communities. Journal of the Fisheries Research Board of Canada
27, 621–651.
Loureau, M. 2000. Are communities saturated? On the relationship between alpha, beta, and gamma diversity.
Ecology Letters 3, 73–76.
Lubchenco, J., Palumbi, S.R., Gaines, S.D. & Andelman, S. 2003. Plugging a hole in the ocean: the emerging
science of marine reserves. Ecological Applications 13, S3–S7.
MacArthur, R.H. 1966. Environmental factors affecting bird species diversity. American Naturalist 98,
387–397.
MacArthur, R.H. 1972. Geographical Ecology: Patterns in the Distribution of Species. New York: Harper &
Row.
Mace, G.M., Gittleman, J.L. & Purvis, A. 2003. Preserving the tree of life. Science 300, 1707–1709.
Mackie, A.S.Y., Oliver, P.G. & Rees, E.L.S. 1995. Benthic biodiversity in the southern Irish Sea. Studies in
marine biodiversity and systematics from the National Museum of Wales. BIOMOR Reports 1, 1–263.
MacNally, R., Bennett, A.F., Brown, G.W., Lumsden, L.F., Yen, A., Hinkley, S., Lillywhite, P. & Ward, D.
2002. How well do ecosystem-based planning units represent different components of biodiversity?
Ecological Applications 12, 900–912.
Magurran, A.E. 2004. Measuring Biological Diversity. Malden Massachusets: Blackwell.
Margalef, R. 1958. Information theory in ecology. General Systems 3:36–71.
Margalef, R. 1995. Information theory and complex ecology. In Complex Ecology: the Part-Whole Relation
in Ecosystem, Patten, B.C. & Jorgensen, S.E. (eds). Englewoods Cliffs: Prentice Hall, 40–50.
Maurer, B.A. 1999. Untangling Ecological Complexity: the Macroscopic Perspective. Chicago: University of
Chicago Press.
May, R.M. 1975. Patterns of species abundance and diversity. In Ecology and Evolution of Communities, M.L.
Cody & J.M. Diamond (eds). Cambridge Massachusetts: Harvard University Press, 81–120.
May, R.M. 1990. Taxonomy as destiny. Nature 347, 129–130.
May, R.M. 1992. Bottoms up for the oceans. Nature 357, 278–279.
169
ROBERT S. CARNEY
McGill, H. 2003. Strong and weak tests of macroecological theory. Oikos 102, 679–685.
Mills, E.L. 1978. Edward Forbes, John Gwyn Jefferys, and the British dredging before the “Challenger”
Expedition. Journal of the Society for the Bibliography of Natural History 8, 507–536.
Murray, J. 1895. A summary of scientific results obtained at the soundings, dredging, and trawling stations
of the HMS Challenger. In Report of the Scientific Results of the H.M.S. Challenger during the years
Years 1873–1876. Edinburgh: Neill and Company.
Narayanaswamy, B.E. Nickell, T.D. & Gage, J.D. 2003. Appropriate levels of taxonomic discrimination in
deep-sea studies: species vs family. Marine Ecology Progress Series 257, 59–68.
National Research Council. 1995. Understanding Marine Biodiversity: A Research Agenda for the Nation.
Committee on Biological Diversity. Washington, D.C.: National Academy Press.
Norse, E.A. (ed.) 1993. Global Marine Biological Diversity: A Strategy for Building Conservation into
Decision Making. Washington, D.C.: Island Press.
Norse, E.A., Rosenbaum, K.L., Wilcove, D.S., Wilcox, B.A., Romme, W.H., Johnston, D.W. & Stout, M.L. 1986.
Conserving Biological Diversity in Our National Forests. Washington, D.C.: The Wilderness Society.
Odum, H.T., Cantlon, J.E. & Kornicker, L.S. 1960. An organizational hierarchy postulate for the interpretation
of species-individual distributions, species entropy, ecosystem evolution, and the meaning of a species-
variety index. Ecology 41, 395–399.
Olsgard, F., Somerfield, P.J. & Carr, M.R. 1998a. Surrogates in marine benthic investigation — which
taxonomic unit to target? Journal of Aquatic Ecosystem Stress and Recovery 7, 25–36.
Olsgard, F., Somerfield, P.J. & Carr, M.R. 1998b. Relationships between taxonomic resolution and data
transformation in analyses of a macrobenthic community along an established pollution gradient.
Paterson, G.L.J. & Lambshead, P.J.D. 1995. Bathymetric patterns of polychaete diversity in the Rockall Trough,
northeast Atlantic. Deep-Sea Research I 42, 1199–1214.
Pearson, D.L. 1994. Selecting indicator taxa for the quantitative assessment of biodiversity. Philosophical
Transactions of the Royal Society of London Series B 345, 75–79.
Peet, R.K. 1974. The measurement of species diversity. Annual Review of Systematics and Ecology 5, 285–307.
Petersen, C.G.J. 1918. The sea bottom and its production of fish food: a survey of the work done in connection
with valuation of the Danish waters from 1883–1917. Reports of the Danish Biological Station 25, 1–62.
Pielou, E.C. 1966. Shannon’s formula as a measure of specific diversity: its use and misuse. American
Pielou, E.C. 1969. An Introduction to Mathematical Ecology. New York: Wiley-Interscience.
Pineda, J. & Caswell, H. 1998. Bathymetric species-diversity patterns and boundary constraints on vertical
range distributions. Deep-Sea Research II 45, 83–101.
Poore, G.C.B. & Wilson, G.D.F. 1993. Marine species richness. Nature 361, 597–598.
Posadas, P., Miranda-Esquivel, D.R. & Crisci, J.V. 2001. Using phylogenetic diversity measures to set priorities
in conservation: an example from southern South America. Conservation Biology 15, 1325–1334.
Preston, F.W. 1948. The commoness and rarity of species. Ecology 29, 254–283.
Pueyo, S. 2006. Self-similarity in species-area relationship and in species abundance distributions. Oikos 112,
156–162.
Purvis, A. & Hector, A. 2000. Getting the measure of biodiversity. Nature 405, 212–219.
Quijón, P.A. & Snelgrove, P.V.R. 2006. The use of coarser taxonomic resolution in studies of predation on
marine sedimentary fauna. Journal of Experimental Biology and Ecology 330, 159–168.
Rex, M.A., Crame, J.A., Stuart, C.T. & Clarke, A. 2004. Large-scale biogeographic patterns in marine mollusks:
a confluence of history and productivity. Ecology 86, 2288–2297.
Rex, M.A., Etter, R.J. & Stuart, C.T. 1997. Large-scale patterns of species diversity in the deep-sea benthos.
In Marine Biodiversity: Patterns and Processes, R.F.G. Ormond et al. (eds). Cambridge: Cambridge
University Press, 94–121.
Rex, M.A., McClain, C.R., Johnson, N.A., Etter, R.J., Allen, J.A., Bouchet, P. & Waren, A. 2005. A source-
sink hypothesis for abyssal biodiversity. American Naturalist 165, 163–178.
Rex, M.A., Stuart, C.T. & Coyne, G. 2000. Latitudinal gradients of species richness in the deep-sea benthos
of the North Atlantic. Proceedings National Academy of Sciences of the United States of America 97,
4082–4085.
170
Ricklefs, R.E. 2004. A comprehensive framework for global patterns in biodiversity. Ecology Letters 7, 1–15.
Rosenberg, M.S., Adams, D.C. & Gurevitch, J. 2000. MetaWin. Version 2.0. Statistical Software for Meta-
Analysis. Sunderland, Massachusets: Sinauer Associates.
Rosenzweig, M.L. 1995. Species Diversity in Space and Time. Cambridge: Cambridge University Press.
Roy, K., Jablonski, D. & Valentine, W.V. 2000. Dissecting latitudinal diversity gradients: functional groups
and clades of marine bivalves. Proceedings Royal Society London Series B 267, 293–299.
Rumohr, H., Karakassis, I. & Jensen, J.N. 2001. Estimating species richness, abundance and diversity with
70 macrobentic replicates in the western Baltic Sea. Marine Ecology Progress Series 214, 103–110.
Russell, R., Wood, S.A., Allison, G. & Menge, B.A. 2006. Scale, environment, and trophic status: the context
dependency of community saturation. American Naturalist 167, E158–E170.
Salas, F., Patrico, J., Marcos, C., Pardal, M.A., Perez-Ruzafa, A. & Marques, J.C. 2006. Are taxonomic
distinctness measures compliant to other ecological indicators in assessing ecological status? Marine
Pollution Bulletin 52, 816–829.
Sanders, H.L. 1960. Benthic studies in Buzzards Bay. III. The structure of the soft-bottom community.
Sanders, H.L. 1968. Marine benthic diversity. American Naturalist 102, 243–282.
Sanders, H.L., Hessler, R.R. & Hampson, G.R. 1965. An introduction to the study of the deep-sea benthic
faunal assemblages along the Gay Head-Bermuda transect. Deep-Sea Research 12, 845–867.
Shannon, C.E. 1948. A mathematical theory of communication. The Bell System Technical Journal 27,
379–423; 623–656.
Shelford, V.E. 1935. General survey of the communities — their extent and dynamics. The major communities.
Ecological Monographs 5, 251–292.
Simberloff, D.S. 1972. Properties of the rarefaction diversity measurement. American Naturalist 106, 414–418.
Simpson, E.H. 1949. Measurement of diversity. Nature 163, 688.
Slavin, R.E. 1986. Best-evidence synthesis: an alternative to meta-analytic and traditional reviews. Educational
Researcher 15, 5–11.
Smith, J.E. 1932. The shell gravel deposits and the infauna of the Eddystone grounds. Journal of the Marine
Biological Association United Kingdom 18, 243–78.
Sneath, P.H.A. & Sokal, R.R. 1973. Numerical Taxonomy. San Francisco: Freeman.
Snelgrove, P.V.R. & Smith, C.R. 2002. A riot of species in an environmental calm: the paradox of the species-
rich deep-sea floor. Oceanography and Marine Biology: An Annual Review 40, 311–342.
Somerfield, P.J., Olsgard, F. & Carr, M.R. 1997. A further examination of two new taxonomic distinctness
measures. Marine Ecology Progress Series 154, 303–306.
Spärck, R. 1935. On the importance of quantitative investigation of the sea bottom in marine biology. Journal
du Conseil 10, 3–19.
Stephen, A.C. 1928. Notes on the quantitative distribution of molluscs and polychaetes in certain areas on the
Scottish coast. Proceedings of the Royal Society Edinburgh 21, 205–216.
Stephen, A.C. 1934. Studies on the Scottish marine fauna: quantitative distributions of the echinoderms and
the naturalistic divisions of the North Sea. Transactions of the Royal Society Edinburgh 57, 777–787.
Stuxburg, A. 1883. Researches on the deep-sea fauna from a zoogeographic point of view. Nature 23, 394–397.
Su, J.C., Debinski, D.M., Jakubauskas, M.E. & Kindscher, K. 2004. Beyond species richness: community
similarity as a measure of cross-taxon congruence for coarse-filter conservation. Conservation Biology
18, 167–173.
Terborgh, J.W. & Faaborg, J. 1980. Saturation of bird communities in the West Indies. American Naturalist
116, 178–195.
Thorson, G. 1955. Modern aspects of marine level-bottom animal communities. Journal of Marine Research
14, 387–397.
Thorson, G. 1957. Bottom communities. In Treatise on Marine Ecology and Paleoecology, J.W. Hedgepeth
(ed.). The Geological Society of America Memoir 67, 461–534.
Thrush, S.F., Lawrie, S.M., Hewitt, J.E. & Cummings, V.J. 1999. The problem of scale: uncertainties and
implications for soft bottom marine communities and their assessment of human impacts. In Bio-
geochemical Cycling and Sediment Ecology NATP AEW, J.S. Gray et al. (eds). Dordrecht: Kluwer
Scientific, 195–210.
171
ROBERT S. CARNEY
Tyler, P.A. (ed.) 2003. Ecosystems of the Deep Oceans. Amsterdam: Elsevier.
Ugland, K.I. & Gray, J.S. 1982. Lognormal distributions and the concept of community equilibrium. Oikos
39, 171–178.
Ugland, K.I. & Gray, J.S. 1983. Reanalysis of Caswell neutral models. Ecology 64, 603–605.
Ugland, K.I. & Gray, J.S. 2003. The species accumulation curve and estimation of species richness. Journal
of Animal Ecology 72, 888–897.
Ugland, K.I. & Gray, J.S. 2004. Estimation of species richness: analysis of the methods developed by Chao
and Karakassis. Marine Ecology Progress Series 284, 1–8.
Ugland, K.I., Gray, J.S. & Ellingsen, K.E. 2003. The species-accumulation curve and estimation of species
richness. Journal of Animal Ecology 72, 888–897.
Ugland, K.I., Gray, J.S. & Lambshead, P.J.D. 2005. Species accumulation curves analyzed by a class of null
models discovered by Arrhenius. Oikos 108, 263–274.
United Nations Conference on Environment and Development. 1992. Convention on Biological Diversity.
Secretariat at the Convention on Biological Diversity. United Nations Environmental Program. Avail-
able HTTP: http://www.biodiv.org/convention/articles.shtml?a=cbd-02.html. Acessed 11 February
2007.
Valdovinos, C., Navanete, S.S. & Marquet, P.A. 2003. Mollusk species diversity in the Southeast Pacific: why
are there more species towards the pole? Ecography 26, 139–144.
Veech, J.A., Summerville, K.S., Crist, T.O. & Gering, J.C. 2002. The additive partitioning of species diversity:
recent revival of an old idea. Oikos 99, 3–9.
Warwick, R.M. 1988. The level of taxonomic discrimination required to detect pollution effects on marine
benthic communities. Marine Pollution Bulletin 19, 259–268.
Warwick, R.M. & Clarke, K.R. 1995. New ‘biodiversity’ measures reveal a decrease in taxonomic distinctness
with increasing stress. Marine Ecology Progress Series 129, 301–305.
Warwick, R.M. & Clarke, K.R. 1998. A taxonomic distinctness index and its statistical properties. Journal of
Applied Ecology 35, 523–531.
Warwick, R.M. & Clarke, K.R. 2001. Practical measures of marine biodiversity based on relatedness of species.
Warwick, R.M., Dashfield, S.L. & Somerfield, P.J. 2006. The integral structure of a benthic infaunal assem-
blage. Journal of Experimental Marine Biology and Ecology 330, 12–18.
Whitakker, R.H. 1960. Vegetation of the Siskiyou Mountains Oregon and California. Ecological Monographs
30, 279–338.
Whitakker, R.H. 1972. Evolution and measurement of species diversity. Taxon 21, 213–251.
Whitfield, J. 2002. Ecology: neutrality versus the niche. Nature 417, 480–481.
Widdicombe, S. & Austen, M.C. 2001. The interaction between physical disturbance and organic enrichment:
an important element in structuring benthic communities. Limnology and Oceanography 46,
1720–1733.
Weiner, N. 1948. Cybernetics, or Control and Communication in the Animal and the Machine. Cambridge,
Massachusetts: The Technology Press.
Williams, P.H. & Gaston, K.J. 1994. Measuring more of biodiversity — can higher-taxon richness predict
wholesale species richness? Biological Conservation 67, 211–217.
Williamson, M. & Gaston, K.J. 2005. The lognormal distribution is not an applicable null hypothesis for the
species-abundance distribution. Journal of Animal Ecology 74, 409–422.
Wilson, G.D.F. 1998. Historical influences on deep-sea isopod diversity in the Atlantic Ocean. Deep-Sea
Research Part II: Topical Studies in Oceanography 45, 279–301.
Wilson, M. & Shumida, A. 1984. Measuring beta diversity with presence-absence data. Journal of Ecology
72, 1055–1064.
Witman, J.D., Etter, R.J. & Smith, F. 2004. The relationship between regional and local species diversity in
marine benthic communities: a global perspective. Proceedings National Academy of Sciences U.S.A.
1001, 15,664–15,669.
Yule, G.U. 1944. Statistical Study of Literary Vocabulary. Cambridge: Cambridge University Press.
Zenkevitch, L. 1963. Biology of the Seas of the USSR. London: Wiley Interscience.
Zipf, G.K. 1935. Psycho-Biology of Languages. Boston: Houghton-Mifflin.
172

Taylor & Francis
CORAL REEFS OF THE ANDAMAN SEA —

AN INTEGRATED PERSPECTIVE
BARBARA E. BROWN
School of Biology, University of Newcastle, Newcastle upon Tyne NE1 7RU, U.K.
E-mail: ProfBarbaraBrown@aol.com
Abstract The Andaman Sea lies on the eastern edge of the Indian Ocean, bordered to the west
by an arc of islands stretching from northern Sumatra to the Irrawaddy delta. Fringing reefs are
abundant in the Andaman and Nicobar Islands (India), Mergui Archipelago (Myanmar), west coasts
of Thailand and Malaysia and northwest Sumatra (Indonesia). Most have never been visited by
scientists because of political constraints; consequently the region is one of the least studied coral
reef areas in the world. Many inshore reefs are intertidal and occur in turbid settings, while off-
shore reefs exist in clearer waters. Regardless of physical rigours, reefs generally display high cover
and high coral diversity. The Andaman Sea has a complex geological history, a varied seafloor
topography, a highly dynamic oceanography and a large tidal range (2–5 m) coupled with periodic
sea-level depressions. It is also a major sink for sediments from the Irrawaddy, the world’s fifth
largest river in terms of suspended sediment load. Human-made influences are limited; sedimen-
tation from land reclamation and dredging are a principal negative factor though rising sea tem-
peratures present a major threat. Natural damage results from aerial exposure on low tides, negative
sea-level anomalies, earthquakes and tsunamis. The dynamic nature of the Andaman Sea and the
in-built stress resistance of many shallow water corals could result in the region being an important
‘refuge’ during an era of global warming.
Introduction
Charles Darwin first described the extent of reef building in the eastern Indian Ocean in his book
The Structure and Distribution of Coral Reefs published in 1842. He wrote: “The coast of Malacca,
Tanasserim, and the coasts northward, appear in the greater part to be low and muddy: where reefs
occur, as in parts of the Malacca Straits, and near Singapore, they are of the fringing kind; but the
water is so shoal, that I have not coloured them” referring latterly to his shading of reef types on
his map of worldwide reef distribution (his Appendix, p. 226). Similarly he dismissed reefs of the
Andamans where recent publications of the day led him to doubt their existence (citing Asiatic
Researches 4, 402). He acknowledged fringing reefs in the Nicobars which extended between 200
and 300 yards (185–277 m) from the shore, while for west Sumatra he commented on numerous
reefs and banks.
Since publication of Darwin’s book there have been several European-based expeditions to the
region (Rao & Griffiths 1998) with much significant marine biology being done by the British
surgeon-naturalists Alcock and Sewell. By 1933, some 90 yr after Darwin’s first mention of reefs
in the region, it was recognised that extensive fringing reefs were indeed present throughout the
Andaman Sea. They are particularly well developed in the Andaman and Nicobars, along the
coastline of Myanmar and Thailand and the northwest tip of Sumatra and to a limited extent around
the off-shore islands of Malaysia and along the Malacca Straits (Figure 1).
173
BARBARA E. BROWN
95° East 100°
Irrawaddy
Delta
MYANMAR
15° 15°
Andaman
Islands
Mergui
ANDAMAN SEA
South
10° Ten Degree Channel 10°
THAILAND
Nicobar Islands Phuket
The Great Channel
Fringing Reef
5° 0 5°
200 km SUMATRA
95° 100°
Figure 1 Distribution of fringing reefs in Andaman Sea, based on information held in Reef Base (www.reefbase.
org). Bold lines along shorelines represent fringing reefs.
Unfortunately up to the present day the Andaman Sea has never been viewed as an integrated
entity and so the full biological significance of its fringing coral reefs has never been recognised.
One reason for this is that major international initiatives under the aegis of the United Nations
Environment Programme (UNEP: Regional Seas Programme), the International Union for Conser-
vation of Nature and Natural Resources (IUCN) and the Global Coral Reef Monitoring Network
(GCRMN) have followed political boundaries. Such demarcations divide the Andaman Sea in half
partitioning the Andaman and Nicobar Islands to South Asia programmes and Sumatra, Thailand
and Myanmar to southeast Asia. Yet, as this review will explore, the reefs within the region
experience many common and unique characteristics.
Geological history, seafloor topography and related coral

biogeography of the Andaman Sea
The geology and plate tectonics of the Andaman Sea are extremely complex and the geological
history of the region is inextricably linked to the tectonics and geological history of Myanmar, the
Andaman and Nicobar Islands, Sumatra and the Malay peninsula (Curray 2005). Figure 2 outlines
the tectonic setting of the Andaman Sea, its islands and the mainland of southeast Asia. The
Andaman Sea extends some 1200 km from Myanmar to Sumatra and 650 km from the Malay
174
CORAL REEFS OF THE ANDAMAN SEA — AN INTEGRATED PERSPECTIVE
East
90° 95° 100°
20° 20°
Fault
EURASIA PLATE
Sagaing
MYANMAR
INDIA PLATE
THAILAND
15° 15°
BURMA
MICROPLATE
Andaman Islands
Andaman Basin
South
10° 10°
ANDAMAN SEA
SUNDA PLATE
Nicobar Islands Phuket

St
ra
it
f
o
Banda Aceh M
al
ac
5° ca 5°
Volcanoes
Su
m
Fault Types at
ra
Thrust U
S
N Simeulue
Strike slip DA
Normal TR
EN
Other CH
km
0 200 400
0° 0°
90° 95° 100°
Figure 2 Tectonic structure of Andaman Sea. The naming of faults follows Pubellier et al. (2003).
peninsula to the Andaman and Nicobar Islands. Major tectonic features affecting the present appear-
ance of the area include collision between the Indian and Eurasian plates, coupling and decoupling
of platelets, crustal movement along fault lines, rotation of continental blocks and opening of
marginal basins, such as the Andaman Sea (Khan & Chakraborty 2005). Where the two major
175
BARBARA E. BROWN
plates meet, the oceanic (Indian) plate subducts beneath the continental Eurasia plate, causing
clockwise rotation of the subduction zone and an increase in obliquity of the convergence (Curray
2005). The Indian plate begins its descent into the Earth’s mantle at the Sunda trench, which is a
surface expression of the plate interface between the Australian and Indian plates situated to the
southwest of the trench, and the Burma and Sunda plates, located to the northeast. Spencer (in press)
describes the convergence as being partitioned into two components comprising both trench-normal
subduction and forces parallel to the trench which generate strike-slip motions along major fault
systems. As a result a sliver plate, the Burma plate, has sheared off parallel to the subduction zone
and is located between the convergent margin to the west and the great fault systems (the Sumatra
fault, West Andaman Fault, and Sagaing Fault to the east).
In terms of timing of this tectonic activity the region has been significantly affected by at least
three major geological events over the last 50 million yr. According to reconstructions by Hall
(1998) the first major event occurred 50 million yr ago (Mya) with changes in the plate boundaries
from the collision of India with Eurasia. At this time resultant mountain building led to major
changes in both habitats and climates and was accompanied by significant changes in drainage
systems. Huge volumes of sediment moved south from central Asia to the sedimentary basins of
the Sunda Shelf. Approximately 25 Mya plate boundaries and motions changed once more due to
the collision between the north Australian margin and arcs to the north. This tectonic event was
probably very important, in terms of biogeography, because it led to new links between Australia
and Southeast Asia across areas which included many shallow marine habitats. Approximately
5 Mya the positions and boundaries of the tectonic plates changed again with the rotation of north
Sumatra and the partial coupling of the Burmese plate with the northeast-moving India plate. The
Burma plate began to move north on the Sagaing fault, leading to the stretching of the Sunda
continental margin north of Sumatra and to ocean crust formation in the Andaman Sea. While many
workers have concluded that the Andaman Sea showed active spreading only during the past 4–5
Mya, others argue that spreading took place in two phases, one in the middle Miocene (~11 Mya)
and another in the late Miocene–early Pliocene (4–5 Mya) (see Khan & Chakraborty 2005 for
review). In terms of rate of spreading Guzman-Speziale & Ni (1993) estimated the spreading rate
to be 3–5 cm yr−1 but more recent estimates (Curray 2005) put the value at 12 mm yr−1.
Such physical upheaval over the last 50 million yr in this part of Southeast Asia is reflected in
an active seismic history with records of earthquakes, uplift and subsidence in the Andaman and
Nicobar Islands from the mid-1800s onward (Bilham 2005, Bilham et al. 2005). Large earthquakes
have been recorded in these islands in 1847, 1881 and 1941 and most recently in nearby Sumatra
in 2004 and 2005 and the Nicobars in 2005. Geological evidence for vertical motion of the islands
dates back to Oldham (1884) who observed extensive uplifted marine terraces (2–2.6 m above sea
level) throughout the coast of South Andaman Island (Bilham et al. 2005). The Andaman and
Nicobar Island chain represent the peaks of a prominent ocean rise extending from eastern Myanmar
to Sumatra. Geology of the islands has been reviewed by Madhaven et al. (1997), Pal et al. (2003)
and Curray (2005). In brief, the assemblage of rocks found on the islands include the ophiolite
complex of the late Cretaceous era, comprising a periotite, mafic and acidic suite of rocks; basaltic
pillow lavas of the Upper Cretaceous to Palaeocene eras; massive silty stones, thinly bedded chert,
pale yellow- and green-coloured limestone of the Palaeocene-Oligocene era and shelly limestones,
fossil coral reefs and beach sand of recent age (Madhaven et al. 1977). The only active volcano in
the Andaman Basin is Barren Island in the Andamans which last erupted in 1803 (Rodolfo 1969).
Recent geological history suggests that the Pliocene-Pleistocene glaciations, which resulted in
pronounced regressions and alternating transgressions of sea water, have also had an effect upon
the region. About 18,000 yr ago sea level was about 120 m lower than today (Haneburth et al.
2000). In the Indo-Malaysian region land replaced much of the seas and bays and created an almost-
complete barrier between the Indian and Pacific Oceans — the emergence of this area, known as
176
the northern Sunda Shelf, saw the extermination of extensive fringing coral reefs now evident as
fossils along relicts of the Malaysian coastline (Tija 1980). Subsequent sea-level rise about 14,000 yr
ago was rapid (16 m in 300 yr) due to a major melting event in polar regions (Haneburth et al.
2000). A sea-level curve derived from Singapore (Hesp et al. 1998) indicates that present sea level
was reached between 6500 and 7000 yr ago; it then rose to +3 m before falling to present mean
sea level about 3000 yr ago. The Singapore sea-level curve is supported by work of Tudhope &
Scoffin (1994) and Scoffin & Le Tissier (1998) for Phuket, Thailand, in the Andaman Sea which
showed that reef growth began here about 6000 yr ago when the low spring tide level was at least
1 m above its present height.
The geological history of the region ultimately shapes the topography of the Andaman Sea
floor (Figure 3). The major features have been summarised by Rodolfo (1969) and Curray (2005).
The eastern portion of this marginal sea is dominated by the Malay continental margin, a 250-km
wide shelf. An inner shelf, less than 100 m deep, encloses the Mergui Archipelago off Myanmar;
the shelf gradually narrowing toward the south where it terminates in a minor slope with 100 m
relief. Off Phuket the inner shelf is only 35 km wide, merging to the south and east with the Sunda
Shelf and the Malacca Strait. Another major feature is the continental slope off the Malay peninsula
with an outer shelf break that occurs at increasing depth southward to a maximum at 7°N latitude;
pinnacles (200 m relief) mark the shelf break. The continental slope ends abruptly in a deep terrace
2435 m deep between the Sewell Rise and Martaban Canyon. This deep terrace slopes gently
westward for 60 km to a depth of 2670 m where the sea floor drops steeply to 3075 m in the
Central Andaman Trough. The Andaman-Nicobar Ridge is another significant feature which bounds
the western portion of the Andaman Sea. The western coastlines of the Andaman and Nicobar
Islands show little indentation and coastal plains are found only on these shores. The eastern shores
are more indented and steep and in many places coral reefs and beaches are raised as high as 20 m
above sea level (Sewell 1925). Major channels cross the Andaman-Nicobar Ridge divide the islands
into four groups which are fringed by coral shelves 10–50 km wide west of the islands and less
than 10 km wide on the east. It has been reported by Alcock (1902), Sewell (1925, 1935) and
Rodolfo (1969) that the Andaman Islands are parallelled 22 km off their western coasts by a series
of coral banks that have been described as a discontinuous barrier reef but these reports remain
unconfirmed. The major channels between the islands include the Great Passage between Great
Nicobar and Sumatra and the Ten Degree Channel between Car Nicobar and Little Andaman. To
the west and northeast, Bay of Bengal and Andaman Sea waters interconnect through narrow
channels 1000–2000 m deep.
Coral biogeography of the region is closely linked to the geological and tectonic history of the
area. Some of the earliest global records of scleractinian corals are found in the Andaman Sea
region (Wilson & Rosen 1998) with upper Triassic (~180 Mya) scleractinian coral recorded on the
Myanmar/Thailand border and in the Indonesian Archipelago (Stanley 1988). Other records include
Upper Jurassic (~140 Mya) known from Sumatra, Myanmar and Thailand (Beauvais 1983) and a
few corals from the Upper Cretaceous (~80 Mya) in northern Sumatra (Wilson & Rosen 1998).
Generally corals were most common throughout Southeast Asia in the Upper Triassic and Upper
Jurassic, reflecting times when corals flourished globally. Wilson & Rosen (1998) concluded that
diversity of recent corals in the region has ultimately been controlled by plate tectonics. At the
beginning of the Tertiary (~70 Mya) Australia was separated from mainland Southeast Asia by
~3000 km of ocean creating an ‘Indo-Pacific gateway’ which narrowed over the next 70 Mya as
Australia moved northward. This fact combined with the emergence of new islands and shallow-
water carbonate areas gave rise to potential for exchange of coral larvae with other regions and the
establishment of coral communities. At the same time, during India’s drift northward suitable
habitats for coral development may have occurred on narrow shelves with limited sediment input
in the eastern Indian Ocean. Wilson & Rosen (1998) commented that the large deltas of the Irrawaddy
177
BARBARA E. BROWN
East
95° 100°
n
anyo
15° 15°
MYANMAR
an C
tab
Mar
2000 m
Inner Shelf
Alcock Rise
0m
300
2000 m
Central
500 m
South Andaman
Trough
10° 10°
200 m
Sewell Rise
THAILAND
2000 m
5° 5°
SUMATRA
95° 100°
Figure 3 Seafloor topography of Andaman Sea. Bathymetry reproduced from GEBCO Sheet G.08 compiled
by R.L. Fisher of Scripps Institution of Oceanography and extracted from GEBCO Digital Atlas published
by British Oceanographic Data Centre for IOC and IHO, 2003.
and Ganges currently inhibit coral growth in their immediate vicinity, forming coastal barriers to
biogeographical exchange of many shallow-water organisms in the northern Andaman Sea.
The Pliocene-Pleistocene glaciations and the exposure of the Sunda Shelf would also have
significant implications for coral biogeography (Potts 1983, Myers 1991). The Andaman Sea was
probably bounded at this time by the Andaman and Nicobar Island bridges which may have
promoted speciation and endemism in the Andaman Sea (Satapoomin 2002). Several endemic reef
fishes have been identified (McManus 1985, Satapoomin 2002) though no corals specific to the
Andaman Sea have yet been described. Two Acropora species are endemic to India/Sri Lanka and
the Andaman Sea (Wallace & Muir 2005). The authors warn that present identifications under-
represent the overall species composition of the region and, as a result, potential endemicity.
178
A recurrent feature of papers dealing with the biogeography of the region is that it has been an
area where the scope to develop diverse communities has been controlled by the way suitable habitats
have been continually created, reorganised and lost through tectonic movements (Wilson & Rosen,
1998) and natural events such as volcanism, sediment influx and storm damage (McManus 1985).
Physical influences affecting coral reefs

Oceanography
Circulation of the Indian Ocean is strongly influenced by the seasonal reversal of the monsoon
winds and their effect on ocean currents in the Northern Hemisphere as described in Tomczak &
Godfrey (1994). When the northeast monsoon is established the North Equatorial Current runs as
a narrow current of about 0.3 m s−1 from the Malacca straits to southern Sri Lanka passing through
the Andaman Sea en route. The transition from northeast to southwest monsoon is characterised
by the easterly moving, intense Indian Equatorial Jet first described by Wyrtki (1973) with velocities
of 0.7 m s−1 or more. When the southwest monsoon is fully established, during July–September,
the northern Indian Ocean is dominated by the eastern flow of the South West Monsoon Current
which enters the Andaman Sea via the Bay of Bengal. The transition before the onset of the northeast
monsoon is also characterised by the equatorial jet which concentrates eastward flow in a 600-km
wide band along the equator. It reaches its peak in November with velocities of 1.0–1.3 m s−1 and
disappears in early January when the cycle is repeated (Tomczak & Godfrey 1994).
A comprehensive review of oceanography of the Bay of Bengal and Andaman Sea is provided
by Varkey et al. (1996) and combines actual observations by Varkey (1986) with results of a
simulation model driven by climatological monthly mean winds (Potemra et al. 1991). Potemra’s
model was a four-layer isopycnal model that used the 200-m contour as the land boundary. Both
Varkey et al. (1996) and Potemra et al. (1991) described surface circulation in the Andaman Sea
as a double gyre with anticlockwise flow during the northeast monsoon and clockwise flow in the
southwest monsoon. Earlier work by Soegiarto & Birowo (1975) and Soegiarto (1985) and simu-
lations using the Ocean Circulation and Climate Advanced Model (OCCAM) by P. Hyder (personal
communication) do not reflect these gyres but do broadly agree on the predominant direction of
current flow throughout the year as shown in Figure 4. This figure is probably an overly simplistic
representation of current flow but is the best available at the present time. The earlier model used
by Potemra et al. (1991) suggests surface flow enters the Andaman Sea south of the Nicobar Islands
during the northeast monsoon and exits south of the Andamans. During the southwest monsoon
flow enters from the Bay of Bengal, circulates clockwise and exits via the southern Andaman Sea.
Khokiattiwong (1991) found that off the west coast of Thailand shallow-water currents were
strongly influenced by tidal currents during the northeast monsoon. In later work S. Khokiattiwong
(personal communication) suggested that there are two major water masses influencing flow around
Phuket on the west coast of Thailand. One, a northern water mass, is described as flowing clockwise
from the Bay of Bengal along the Andaman Sea coast of Thailand to Phuket where it meets a
southern water mass flowing from the Malacca Strait northward in a counterclockwise direction
before it flows east to the Indian Ocean. Between the two water masses there is a mixing zone
which shifts further to the north or south of Phuket depending on the effects of the reversing
monsoon. Such interpretations are based on salinity and temperature distributions and measurements
of sea-level height.
The highly dynamic nature of the Andaman Sea is reflected in current flow at depth (>200 m)
where Potemra et al. (1991) suggested that flow changes direction three times a year. In January,
February and March flow is clockwise; between April and July it is anticlockwise; from August to
October it becomes clockwise before reverting to anticlockwise in November and December.
179
BARBARA E. BROWN
East
95° 100°
Myanmar
15° 15°
Andaman
Islands
South
10° 10°
Thailand
Nicobar
Islands
5° 5°
Sumatra
N.E. Monsoon
25 50 75 cm/sec
Myanmar
15° 15°
Andaman
Islands
South
10° 10°
Thailand
Nicobar
Islands
5° 5°
Sumatra
S.W. Monsoon
95° 100°
Figure 4 Predominant surface currents in the Andaman Sea in the northeast monsoon (December–May) and
the southwest monsoon (June–November) after Soegiarto & Birowo (1975) and Soegiarto (1985). Arrows
depict current strength. (Data reproduced with the permission of the United Nations Environment Programme.
UNEP Regional Seas Reports and Studies No. 69 (1985).)
180
While precise details of water movements within the Andaman Sea remain speculative other
hydrographic features have received considerable attention. Large-amplitude, long-interval waves,
with associated surface waves (‘rips’), have been observed in the Andaman Basin (Osborne &
Burch 1980, Osborne 1990, Hyder et al. 2005). These internal waves, which are apparently due to
non-linear internal tides (or solitons), have been observed as surface rips in satellite imagery of the
Andaman Sea (Alpers et al. 1997). The waves propagate within subsurface layers of the sea that
are stratified because of temperature and salinity variations and appear to result from tidal interaction
with tidal sills and seamounts, spreading radially from the source. The length of the wave becomes
longer as the waves spread out from the source.
In the southern Andaman Sea the waves occur in packets of 4–10 and propagate several hundred
kilometres before encountering the Thai coastline. The waves are normally rank ordered by ampli-
tude (crest-to-trough distance) with the largest wave leading the rest. In one event, the amplitude
of the foremost wave was estimated to be 60 m with the warm water from above being pushed
down by the internal soliton by 60 m (Alpers et al. 1997). Bands of surface rips accompany the
internal waves; these bands are 600–1200 m wide, stretch 10–100 km across the sea surface and
can be observed in satellite photographs. In the northern Andaman Sea the waves only occurred on
spring tides when the tidal range exceeded 1.5 m and the probability of their occurrence increased
with tidal range (Hyder et al. 2005).
Internal waves represent a significant mechanism for the transport of momentum and energy
within the ocean (Osborne & Burch 1980) and possibly lead to increased mixing, perturbation of
temperature and salinity gradients and potential increases in primary and secondary production in
coastal waters (Nielsen et al. 2004). In addition to internal waves, a longer-period internal oscillation
has been observed in the northern Andaman Sea resulting in a downward perturbation of the
pycnocline (the density gradient caused by the thermocline) by 80 m in a survey period between
January and April 1998 (P. Hyder personal communication). These observations have been attributed
to eastward-propagating Kelvin-like waves which propagate parallel to the continental slope. Such
waves also have considerable implications for mixing processes in the Andaman Sea.
Upwelling is another significant feature of the Andaman Sea. Yesaki & Jantarapagdee (1981)
suggested that coastal upwelling on the west coast of Thailand is a recurrent phenomenon generated
by the monsoon system. In recent work Nielsen et al. (2004) argued that the shelf area of the
Andaman Sea has the potential of being a productivity ‘hot spot’ with stratified water meeting
mixed coastal water in combination with very dynamic oceanography. While high production was
not noted at the shelf break in this study the site of particularly high production was at the mid-
shelf front where breaking of the shoaling waves introduced cold, nutrient-rich water to the euphotic
zone.
Tidal influences and sea-level fluctuations

Most coral reefs in the Andaman Sea are subject to semi-diurnal tides. Mesotidal areas with a spring
tide range of 1–2 m are found in the Andaman and Nicobar Islands and along the north/northwest
coast of Sumatra while macrotidal ranges are found along the coast of Thailand (~3 m) and the
Mergui Archipelago (>5 m) of Myanmar.
Sea levels in the region show high annual variability (Figure 5) with considerable anomalies
in some years (Dunne & Brown 2001). Both in situ and remote satellite measurements of sea level
along the Thai coastline are well matched over the period shown, apart from two periods of pronounced
positive anomalies reflected in the in situ data in early 1994 and 1997 (which are probably the
result of Kelvin waves moving along inshore areas) but not in the off-shore altimeter record. Large
negative sea-level anomalies of −21 and −27 cm are obvious in late 1994 and late 1997, respectively.
A period of negative sea-level anomalies in 1994–1995 extended over 9 months from May 1994
181
BARBARA E. BROWN
N7.777 E97.05
40
N7.83 E97.07
Ko Taphao Noi
30
Sea level anomalies (cm) 20
10
−10
−20
−30
−40
93 94 95 96 97 98 99
19 19 19 19 19 19 19
Figure 5 Sea-level anomalies for Ko Taphai Noi tide station, Phuket, Thailand (shown as monthly means)
and from TOPEX/POSEIDON satellite altimeter records at two off-shore stations. (From Dunne & Brown
2001. With permission of Springer Science and Business Media.)
to January 1995 whereas the more pronounced negative anomalies of 1997–1998 persisted over
11 months from July 1997 to May 1998. Such negative sea-level anomalies are a regular feature
in the Andaman Sea and Indian Ocean and were first described by Webster et al. (1999). These
authors described the anomalies as the result of ocean-atmosphere-land interactions in the Indian
Ocean which occur primarily as a result of a reversal in sea temperature gradient between east and
west basins of the Indian Ocean. During this process there is substantial warming in the western
basin of the Indian Ocean with higher sea levels, lowered thermocline and reduced upwelling. In
contrast, sea level is depressed in the eastern Indian Ocean and the position of the thermocline is
raised with resultant enhanced upwelling. This phenomenon is known as the Indian Ocean Dipole,
and although Webster et al. (1999) considered the phenomenon internally forced others believe that
the anomalies are externally forced by connection with the Pacific El Niño Southern Oscillation
(ENSO) (Allan et al. 2001). The resultant sea-level depressions may cause significant mortality on
shallow coral reefs throughout the Andaman Sea (Brown & Phongsuwan 2004). Not only are low
tides much lower than normal, involving greater exposure of reef flat organisms for longer periods,
but the times of exposure are also altered. Marine organisms may, as a result, find themselves
exposed at mid-day under maximum solar radiation rather than early morning and evening when
solar radiation is minimal.
Sea temperatures and salinity

Monthly sea temperatures in the Andaman Islands range from 28.1°C in January to a maximum of
29.8°C in May while on the Thai coast temperatures range from 27.8°C in January to a maximum
of 29.5°C in May (Brown et al. 1996). Recent analyses of historical sea temperature data and
contemporary continuous sea-surface measurements in the Andaman Sea at Phuket show an inter-
esting trend for the eastern Indian Ocean (Figure 6) where there has been a significant increase in
sea-surface temperatures over the last 50 yr of at least 0.126°C decade−1 (Brown et al. 1996). This
is consistent with positive trends in the Indian Ocean area demonstrated by others (Hoegh Guldberg
182
1997
1998
1991 1995
30.1°C
30
29
SST °C
28
27
01
51
81
91
71
61
46
56
66
76
86
96
20
19
19
19
19
19
19
19
19
19
19
19
Figure 6 Monthly mean sea-surface temperatures (SST) 1945–2004 from the Meteorological Office Historical
Sea Surface Temperature (MOHSST 6) dataset for Andaman Sea area off Phuket. Regression line for all points
shown (p < 0.001). The 30.1°C line represents an approximate coral bleaching threshold since solar radiation
levels, as well as sea temperature, have a significant part to play in bleaching.
1999) and with current predictions of the Intergovernmental Panel on Climate Change (IPCC 2001).
Such trends have major implications for all marine fauna, particularly many corals which have
been shown to be extremely sensitive to increases in temperature, bleaching (losing colour through
loss of essential algae and/or their pigments) when temperatures exceed their seasonal maxima by
only 1°C.
Salinity measurements in the Andaman Sea are limited. According to Varkey et al. (1996)
surface salinity varies between 32 in the northern Andaman Sea to 33 in the south during the
northeast monsoon. In support of these values Janekarn & Hylleberg (1989) found surface salinities
of between 32.1 and 33.6 in coastal waters of Phuket in the southern Andaman Sea and Desai et al.
(1988) values ranging from 31 to 32.8 in the northern Andaman Sea. Varkey et al. (1996) provided
no salinity data for the Andaman Sea in the southwest monsoon though Tomczak & Godfrey (1994)
reported salinities below 25 in the central Andaman Basin at this time. Desai et al. (1988) described
very low salinities (but give no values) due to river discharge during the southwest monsoon in the
northern Andaman Sea, noting that below the surface salinity increases rapidly with a strong
halocline developing at 10–60 m after the southwest monsoon period.
Sedimentation
Major rivers discharge into the Bay of Bengal, north of the Andaman Sea and directly into the
Andaman Basin and have done so for thousands of years. The Ganges/Brahmaputra delta is the
largest estuarine system in the world with an estimated average annual suspended load of 2,179,000 ×
103 tonnes (mt) and an average discharge of 31.5 × 103 m3 s−1 (Rao & Griffiths 1998). Sediments,
sourced from the Himalayas, are discharged into the Bay of Bengal where they contribute to the
Bengal Fan, which is the world’s largest accumulation of sediments. The Irrawaddy in Myanmar
also makes a significant sedimentary contribution directly into the Andaman Sea with an average
annual suspended load of 300,000 × 103 mt and an average discharge at the mouth of 13.6 ×
103 m3 s−1. Ramaswamy et al. (2004) gave a comprehensive account of the fate of the Irrawaddy
183
BARBARA E. BROWN
discharge providing turbidity profiles that show transport of sediments into the deep Andaman Sea
via the Martaban Canyon.
While sedimentation limits reef growth in the vicinity of these enormous river discharges coral
reefs do flourish in inshore turbid settings on all islands and mainland areas bordering the Andaman
Sea where they receive considerable local drainage from mangrove forests. Indeed these waters,
enriched in particulate and dissolved organic matter, may be of considerable benefit to the corals
of the region as described in later sections of this review.
Biological characteristics of coral reefs in the Andaman Sea

Reef type and coral diversity
Fringing reefs dominate all sites within the Andaman Sea with the only ‘barrier’ reef reported by
Sewell (1935) on the west coast of the Andaman Islands. This observation remains to be confirmed
though Turner et al. (2001) also noted off-shore reef structures on the west side of the Andamans
in a recent survey using satellite imagery. Throughout the Andaman Sea fringing reefs tend to be
better developed on the eastern sides of islands (Phongsuwan & Chansang 1992, Spalding et al.
2001) where they are not exposed to the southwest monsoon influence. In northern Aceh, Sumatra
reef development is mainly on the west side of the mainland and on off-shore islands to the north
(A.H. Baird personal communication).
Extensive intertidal reefs flats are found throughout the region (Figure 7). Sheltered reefs are
dominated by a high cover of Porites species and faviids while those receiving more wave exposure
are dominated by the branching corals Acropora spp., Pocillopora damicornis and Montipora
digitata. Such a distribution pattern associated with wave exposure is common throughout the
whole Indo-Pacific (Rosen 1975) although A.H. Baird (personal communication) noted very little
M. digitata on reefs in Aceh, in northern Sumatra. In the Andamans and Nicobars the width of reef
flats is considerable, ranging from 200 to 500 m (UNEP/IUCN, 1988). Many of these reefs support
significant growth of alcyonarians as well as stony corals. Extensive reef flats may also be found
on the mainland coast of Thailand and occasionally in Aceh. Generally, reefs along the coast of
mainland Thailand are found at depths of 5–15 m whereas those on off-shore islands are found in
depths down to 30 m. A detailed mapping of Thailand’s reefs in the Andaman Sea has been carried
out and the results are published in Chansang et al. (1999). The most abundant species in deeper
water are Porites lutea, Acropora intermedia, A. muricata (formally formosa) and Porites (synaraea)
rus (Phongsuwan & Chansang 1992). These authors also noted that coral communities on off-shore
islands are usually formed of a single species on the reef slope, while those on near-shore islands
comprise mixed species. As a result, diversity is higher at near-shore sites than at off-shore locations.
In addition to fringing reefs, the reefs of some off-shore islands develop principally in bays as
scattered isolated coral heads, clumps and thickets of coral which have a high percentage of living
coral. An array of reef types has been described in the Andamans and Nicobars by Reddiah (1977)
and in the Andamans by Turner et al. (2001). These include channel reefs, which are found on
shorelines between islands where wind and wave influences are less severe with slopes down to
20 m depth, and knoll reefs which occur in channels and patch reefs. Where sediment levels are
high and calm conditions prevail, inshore reefs are dominated by poritid and faviid corals similar
to those described on the west coast of Thailand (Reddiah 1977). In more wave-exposed situations
reef edges are dominated by Acropora, Pocillopora and Montipora. On the northwest coast of
Sumatra the outer reef edges of off-shore islands also support high-energy coral reef associations
of Acropora and Pocillopora (A. Kunzmann personal communication). In the Nicobars, seaward
edges of reefs on the west side of the islands have well-developed spurs and grooves with surge
channels about 1 m deep and 1 m wide and dominated by calcareous red algae (Reddiah 1977).
184
Figure 7 Fringing intertidal reefs in the Andaman Sea. (A) Havelock Island Andamans; reef is dominated by
alcyonarians. (Photograph courtesy Jason Reubens.) (B) Southeast Phuket; reef is dominated by Acropora
aspera, A. pulchra and Montipora ramosa. (Photograph courtesy Barbara Brown.) (C) Paway Island, Mergui
Archipelago, Myanmar; reef is a mixture of massive (Porites sp.) and branching corals. (Photograph courtesy
R.B.S. Sewell.)
Unfortunately very little ecological information is available for the reefs of the Mergui Archipelago
in Myanmar although the southernmost reefs appear to resemble, in terms of coral communities,
those of the Surin Islands in Thai waters to the south. Initial surveys suggest that the majority of
reefs visited in the southern Mergui have >50% cover (Wilkinson 2004).
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BARBARA E. BROWN
Table 1 The number of zooxanthellate corals recorded to date at selected

locations in the Andaman Sea
No. of No. of
Location coral species coral genera Reference
Andaman Islands 57 23 Pillai 1972
Andaman Islands 49 15 Reddiah 1977
Andaman Islands 187 56 Turner et al. 2001
Andaman and Nicobar Islands 110 45 Pillai 1983
Andaman and Nicobar Islands 203 ND Wilkinson 2004
Nicobar Islands 64 19 Reddiah 1977
Nicobar Islands ND 40 Scheer 1971
Mergui Archipelago 51 26 Summarised by Pillai 1972
Mergui Archipelago 65 ND Wilkinson 2004
West coast of Thailand 268 66 Phongsuwan in press
West coast of Thailand 353 69 Turak et al. 2005
Note: ND, not described.
Table 1 summarises the level of diversity of corals throughout the region. The values are very
variable even for a single region and reflect, in the Andaman Islands for example, limited sampling
in earlier studies. In several of the publications cited, corals have been designated variously as
hermatypes or non-hermatypes, zooxanthellate or non-zooxanthellate and scleractinian or non-
scleractinian which makes cross-comparison between studies very difficult In addition, it is likely
that figures given for the Mergui Archipelago and the Nicobars are underestimates because no
comprehensive surveys have been carried out here or on the northwest tip of Sumatra.
While Andaman Sea reefs are some of the most diverse in the Indian Ocean they have previously
been reported as being less diverse than those in the Philippines and Indonesia. Recent surveys in
western Thailand, however, now report 353 coral species (Turak et al. 2005) which brings the
Andaman Sea into the Indo-Pacific ‘coral triangle’ of high biodiversity centred on Indonesia to the
east. Interestingly, a number of these newly recorded coral species were previously known only
from the Pacific.
Despite limited sampling Wallace (1999) noted at least 55 species of Acropora in the Andaman
Sea, a figure which is exceeded in the Indian Ocean only in the ‘eastern Indian Ocean’ which
boasts 71 species and ranks alongside the most diverse areas of the world. Of Acropora species
recorded in the Andaman Sea, 51 are regarded as widespread and are present in 12 of the 29
biogeographic areas described in the Indo-Pacific.
Physiological attributes of reef corals in the Andaman Sea

Early work on zooxanthellae densities in corals suggested that they were remarkably constant,
ranging from 1 × 106 to 2.5 × 106 algae cm−2 (Drew 1972). Subsequent work has shown more
variability particularly in corals from turbid waters such as those which characterise shallow inshore
areas of the Andaman Sea. In reef-flat corals from Phuket, algal densities per square centimetre of
coral tissue ranged from 0. 6 × 107 to 1.4 × 107 in Porites lutea, 0.4 × 107 to 1.8 × 107 in the faviid
Goniastrea retiformis and 0.8 × 107 to 2.6 × 107 in the faviid G. aspera and agaricid Coeloseris
mayeri (Brown et al. 1999). Massive corals from inshore turbid waters around Singapore (B. Goh
personal communication) and Java (Suharsono & Soekarno 1983) have similar high algal densities.
Non-massive reef-slope corals at Phuket also show relatively high algal densities ranging from
186
5 × 106 cm−2 (Psammocora digitifera) to 6 × 106 cm−2 (Mycedium elephantotus) (Satapoomin 1993).
A potential influence on algal density is the nutrient content of surrounding sea water because
experimental studies have shown that the density of algae increases on exposure to elevated nutrient
concentrations (Stambler et al. 1991). The concentration of dissolved nutrients at Phuket is high
all year round due to drainage from extensive mangrove areas to the north and oceanic upwelling
off shore (Janekarn & Hylleberg 1989). Brown et al. (1999) compared levels of dissolved nutrients
at Phuket with other reef sites worldwide and showed that values of nitrate, nitrite and phosphate
recorded at Phuket were high, ranking alongside other inshore locations in the Florida Keys and
Barbados which experienced eutrophication. In addition, particulate load is elevated in inshore
waters, frequently reaching 20–40 mg l−1 (Scoffin et al. 1992, Panutrakul 1996). Particulates are
derived from fine terrigeneous sediments composed of clay minerals. It is likely that these partic-
ulates, coated with bacteria and microalgae, present added potential nutrients to particulate-feeding
corals in the region (Anthony 1999). As a result corals have the potential to benefit from both
autotrophic and heterotrophic feeding.
The algal genus Symbiodinium, which forms a symbiotic relationship with corals, is diverse
and molecular analyses have shown that there is considerable variation at the level of the ribosomal
RNA genes. It comprises two clades, one known as phylotype A and the other which includes
phylotypes B–F (LaJeunesse 2001). Of all the phylotypes so far evaluated phylotype D appears to
be the most thermotolerant (Rowan 2004). Interestingly, phylotype D is present in many shallow-
water corals and zoanthids throughout the Indian Ocean (Goodson 2000, Burnett 2002) whereas
this phylotype is relatively rare on the southern Great Barrier Reef and in the Caribbean (LaJeunesse
et al. 2003). On extensive reef flats of the east coast of Phuket, Thailand phylotype D was identified
in all six species (four genera) sampled. In most cases D was present to the exclusion of all other
phylotypes apart from Acropora pulchra which contained a mixture of C and D. Interestingly, in
Goniastrea aspera, which is a cosmopolitan intertidal reef species throughout the Indo-Pacific,
colonies from the southern Great Barrier Reef contained phylotype C (LaJeunesse et al. 2003)
although only phylotype D has been found in this species at Phuket (Goodson 2000). Since the
rigours of intertidal living are similar on the Great Barrier Reef to those in Thailand it appears that
something other than environmental constraints might be acting to produce this distribution pattern.
Burnett (2002) believes that biogeography may play a major role in the distribution of symbionts
in the zoanthid Palythoa in the Indian Ocean. If this is also true for corals, then the presence in a
wide variety of coral species of the most thermotolerant algal symbiont known could have important
implications for the impacts of global warming.
Another noteworthy feature of corals in the Andaman Sea (in this case specifically massive
corals from the west coast of Thailand) is their rate of skeletal extension, which appears to be
higher than any other recorded in the Indo-Pacific. Buddemeier & Kinzie (1976) described extension
rates for massive corals in optimal settings around the world as 10–15 mm yr−1 with the rate for
Porites spp. being slightly higher. Lough & Barnes (2000) compared the extension rates of P. lutea
from 44 Indo-Pacific reefs (29 on the Great Barrier Reef; 14 in the Hawaiian Archipelago and
1 close to Phuket, Thailand). The Thai study was carried out by Scoffin et al. (1992). Skeletal
extension and calcification was significantly higher in corals from Phuket compared with those
from other Indo-Pacific sites, being two to three times higher than corals from the Great Barrier
Reef and at least a third higher than Hawaiian corals. According to Lough & Barnes (2000) such
differences were strongly linked to sea temperature with the highest sea temperature being found
in the Andaman Sea. In the original study of Thai corals Scoffin et al. (1992) showed that skeletal
extension rates in P. lutea ranged from 1.35 cm yr−1 at off-shore sites to 3.25 cm yr−1 at inshore
locations. The significant difference in extension rates between sites was attributed to wave energy,
with linear extension decreasing along a gradient of increased hydraulic energy. Clearly environ-
mental factors are important in controlling coral skeletogenesis but one factor which has not been
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BARBARA E. BROWN
taken into account above is the nutrient content of sea water. Earlier descriptions in this review of
sites in the Andaman Sea have highlighted the extent of turbid, high-nutrient, sheltered inshore reef
settings which may have a bearing on the remarkably high zooxanthellae densities recorded in
shallow-water corals. If the synthesis of organic matrix (upon which crystals of calcium carbonate
are deposited in the coral skeleton) is a product of the zooxanthellae, as earlier work on calcification
suggests (Johnstone 1980), then the observed rapid skeletal extension of corals in the Andaman
Sea may be a result not only of high sea temperature and reduced wave energy but also of high
nutrients. It is not clear how universal these growth characteristics might be in the Andaman Sea
but it is likely that, because of the similarity of environmental conditions throughout the region, it
is more widespread than the west coast of Thailand.
The high tidal range (2–5 m) at many sites in the Andaman Sea, which favours extensive
intertidal reefs in the region, results in many coral species being exposed to rigorous environmental
conditions during low spring tides. During these periods, corals will be exposed to extremely high
sea temperatures, high solar radiation, and reduced salinities. To survive in these settings the corals
have developed remarkable environmental tolerances to temperature, light, desiccation and salinity.
Protective mechanisms range from behavioural responses which induce rapid tissue retraction
(Brown et al. 2002a) to photoprotective mechanisms such as xanthophyll interconversion and
biochemical defences that include antioxidant enzyme and heat-shock protein production in both
coral host and symbiotic alga (Brown et al. 2002b). Not surprisingly all these mechanisms are
particularly well developed in intertidal corals in the region which are capable of withstanding up
to 3-h aerial exposure at low spring tides. Relatively few assays of such defences have been
monitored worldwide, but in the few studies that have been carried out production of stress responses
by intertidal corals from the Andaman Sea (e.g., Goniastrea aspera) are many-fold higher than
subtidal species such as Montastraea faveolata from the Caribbean when exposed to similar stress
levels (Downs et al. 2000, Brown et al. 2002b). It is interesting to note that although there has been
marked coral bleaching in years with anomalous sea temperatures such as 1991, 1995 and 2003, the
intertidal reefs around Phuket, Thailand, have shown no significant mortality (Brown & Phongsuwan
2004) — a result which is testimony to the fact that these corals are well endowed with effective
environmental defences and have the potential to acclimatise and adapt to varied environments.
Natural and human influences on coral reefs

in the Andaman Sea
The coral reefs of the Andaman Sea have been described as some of the most diverse, extensive
and least disturbed by human intervention in the Indian Ocean (Wallace & Muir 2005) particularly
with respect to the Andamans and Nicobars and Myanmar (Wilkinson 2004). There is generally a
low level of human interference in the region apart from Aceh in Sumatra, where damaging fishing
practices have negatively affected reefs (A.H. Baird personal communication). Thai reef scientists
report very few incidences of damaging fish practices in their waters (U. Satapoomin personal
communication) but highlight the establishment of marine protected areas as a factor which min-
imises such activities. Within the Andaman Sea there are at least 100 nominated marine protected
areas in the Andamans and Nicobars, 2 in Myanmar, 13 in Thailand, 2 on the northwest coast of
peninsular Malaysia and 1 on the northwest tip of Sumatra (Spalding et al. 2001). Enforcement
procedures, however, are not rigorous in the majority of these designated areas.
Despite a low level of human influences shallow reefs are subject to natural disturbances which
include exposure to high solar radiation during aerial exposure, lowered sea level during Indian
Ocean Dipole events, decreased salinity from heavy rain at low tide, tsunami-related damage and
volcanic uplift. There is evidence of marked partial mortality as a result of exposure of faviid
188
colonies to high solar radiation at low tide on reefs in the Andamans, west coast of Thailand and
Malaysia (Brown et al. 1994) and this phenomenon, which is induced by solar bleaching, is probably
widespread throughout the region. Unusually low spring tides also cause coral mortality with sea-
level depressions of 1994 and 1997–1998 (>20 cm experienced over 8–10 months) resulting in
considerable coral mortality on shallow reefs of western Thailand (Brown & Phongsuwan 2004).
Again, these events affect reefs throughout the region, particularly those on the northwest coast of
Sumatra where the sea-level depression of 1997–1998 exceeded 30 cm for several months (Webster
et al. 1999).
This review has already highlighted the fact that earthquakes and resultant tsunamis have been
a recurrent historic feature of the Andaman Sea although the 26 December 2004 earthquake was
unprecedented in scale. Uplifted reef flats (raised by 1–2 m) were reported as a result of the
earthquake on the southwest coast of Simeulue, off the west coast of Sumatra (Sieh 2005) and also
in the Andaman Islands (Searle 2006). Reefs uplifted by the earthquake on the west coast of islands
along the 1000-km arc from Sumatra to the northern most Andamans represent a significant loss
to the gene pool of physiologically resistant corals in this area with many thousands of colonies
now well above the high-tide mark.
A rapid assessment survey of the 700-km coastline of west Thailand (Department of Marine
and Coastal Resources 2005) 4 days after the earthquake revealed very little damage to coral reefs
as a result of tsunami waves. Of the 174 sites visited up to 105 were unaffected or showed very
little damage, with 30 sites displaying low-level damage (11% of coral cover affected). A further
16 sites showed moderate damage (31–50% cover affected) while 23 sites were severely damaged
(>50% coral cover affected). The type of damage fell into three categories: (1) overturned massive
corals, (2) broken branching corals and (3) sedimentation effects. The northernmost coastline and
its off-shore islands were more severely impacted than the south — apart from Phi Phi Island, with
shallow reefs on wave-exposed islands and shorelines that are most vulnerable to wave-induced
damage. Similar results were obtained in Aceh although at this location deeper-living massive
Porites corals, with poor attachment to the substratum, were dislodged while many shallow-living
colonies were unaffected (Baird et al. 2005). At damaged locations in Thailand it is predicted that
recovery will be relatively rapid (5–10 yr) based on monitoring of reefs affected by storm surges
in 1986 (Phongsuwan 1991, Satapoomin et al. 2006).
Reports of human damage to reefs are restricted to the Thai coastline because the majority of
research has been carried out here. They include the effects of tin-smelting operations (Brown &
Holley 1982), tin dredging (Chansang et al. 1992), land reclamation and associated dredging (Brown
et al. 1990, Clarke et al. 1993) and tourism-related activities. While all these perturbations may
cause localised damage, from which coral reefs may recover once the environmental stressor is
removed, global warming represents a much more serious threat to all reefs in the Andaman Sea.
Already a significant increase in sea temperature has been noted in the Andaman Sea over the last
50 yr (Brown et al. 1996), leading to claims that by the late 1990s coral bleaching might be seen
on an annual basis in the region (Hoegh Guldberg 1999). In fact, this has not been the case because
the two most severe bleaching events were in 1991 and 1995 in Thai waters. While the effects of
global warming cannot be underestimated there seems to be little evidence at present to support
so gloomy a prognosis. Not only has there been no annual bleaching of corals but also there has
been limited mortality on both off-shore and inshore reefs. Presently the coral reefs of the Andaman
Sea are in very good condition with inshore corals displaying remarkable physical tolerances. These
in-built tolerances will, however, be severely tested if sea temperatures continue to rise. Rather
than being the first reefs to succumb to global warming inshore reefs in the region may be protected
by both turbid waters and their broad array of environmental defences acquired over centuries. Off-
shore reefs in clearer waters are likely to be more susceptible to global warming, at least initially,
189
BARBARA E. BROWN
because they live in a much more benign environment and have not acquired the same level of
physiological defences as their inshore counterparts.
Conclusions
The coral reefs of the Andaman Sea are not only extensive and largely undisturbed but also boast
a relatively high diversity of corals — the levels of which are likely to be revised upward in the
future because of the present limited surveillance. Such high diversity is probably a consequence
of the complex geological history of the area which has led to habitat disturbance that promoted
increased species diversity over time. Features highlighted in this review, such as the dynamic and
complicated hydrography, the marked tidal ranges and regular sea-level anomalies, will also have
resulted in corals and symbiotic algae developing remarkable environmental tolerances. It is now
abundantly clear that corals thrive in turbid settings such as those in the Andaman Sea and nearby
Java (Bak & Meesters 2000) where they appear to have acclimatised/adapted to living under high
sediment loads. Indeed Potts & Jacobs (2000) suggested that success in turbid habitats allowed
corals to radiate out to more ‘typical’ oceanic habitats. They further proposed that a variety of
turbid inshore habitats have been continuously available through geological time providing eco-
logical and evolutionary continuity as well as refugia for corals during non-optimal periods of reef
growth. The Andaman Sea could well have been such a refuge in the past and indeed may act as
such in the future if sea temperatures continue to rise.
Acknowledgements
I acknowledge the support of staff at Phuket Marine Biological Center over the last 26 years.
Thanks are given to the Natural Environment Research Council (NERC), United Kingdom, the
Leverhulme Trust, United Kingdom, the Royal Society, United Kingdom, and the U.K. Department
for International Development for research support. Thanks also to Peter Hunter at the National
Oceanographic Centre, Southampton, for compilation of Figure 3 and to Dr. Patrick Hyder of the
Meteorological Office for discussion of the oceanography of the Andaman Sea.
References
Alcock, A. 1902. A Naturalist in Indian Seas. London: John Murray.
Allan, R., Chambers, D., Drosdowsky, W., Hendon, H., Latif, M., Nicholls, N., Smith, I., Stone, R. & Tourre,
Y. 2001. Is there an Indian Ocean dipole, and is it independent of the El Nino-Southern Oscillation?
CLIVAR Exchanges 6, 18–21.
Alpers, W., Wang Chen, H. & Hock, L. 1997. Observation of internal waves in the Andaman Sea by ERS
SAR. International Geosciences and Remote Sensing Symposium, Singapore 4, 03–08.
Anthony, K.R.N. 1999. Coral suspension feeding on fine particulate matter. Journal of Experimental Marine
Baird, A.H., Campbell, S.J., Anggoro, A.W., Ardiwijaya, R.L., Fadii, N., Herdiana, Y., Kartawijaya, T.,
Mahyiddin, D., Mukminin, A., Pardede, S.T., Ptrachett, M.S., Rudi, E. & Siregar, A.M. 2005. Acehnese
reefs in the wake of the Asian tsunami. Current Biology 15, 1926–1930.
Bak, R.P.M. & Meesters, E.H. 2000. Acclimatization/adaptation in a marginal environment. Proceedings of
the Ninth International Coral Reef Symposium, Bali, Indonesia 1, 265–271.
Beauvais, L. 1983. Jurassic Cnidaria from the Philippines and Sumatra. Economic and social commission for
Asia and the Pacific. Committee for Co-Ordination Joint Prospecting for Mineral Resources in Asian
Offshore Areas (CCOP) Technical Bulletin 16, 39–67.
190
Bilham, R. 2005. A flying start, then a slow slip. Science 308, 1126–1127.
Bilham, R., Engdhal, E.R., Feld, N. & Satyabalam, S.P. 2005. Partial and complete rupture of the Indo-
Andaman plate boundary 1847–2004. Seisomological Research Letters 76, 299–311.
Brown, B.E., Downs, C.A., Dunne, R.P. & Gibb, S.W. 2002a. Preliminary evidence for tissue retraction as a
factor in photoprotection of corals incapable of xanthophyll cycling. Journal of Experimental Marine
Brown, B.E., Downs, C.A., Dunne, R.P. & Gibb, S.W. 2002b. Exploring the basis of thermotolerance in the
reef coral Goniastrea aspera. Marine Ecology Progress Series 242, 119–129.
Brown, B.E., Dunne, R.P., Ambarsari, I., Le Tissier, M.D.A. & Satapoomin, U. 1999. Seasonal fluctuations
in environmental factors and their influence on photophysiological parameters in four Indo-Pacific
coral species from the Andaman Sea, Indian Ocean. Marine Ecology Progress Series 191, 53–69.
Brown, B.E., Dunne, R.P. & Chansang, H. 1996. Coral bleaching relative to elevated seawater temperature in
the Andaman Sea (Indian Ocean) over the last 50 years. Coral Reefs 15, 151–152.
Brown, B.E., Dunne, R.P., Scoffin, T.P. & Le Tissier, M.D.A. 1994. Solar damage in intertidal corals. Marine
Brown, B.E. & Holley, M.C. 1982. Metal levels associated with tin dredging and smelting and their effect
upon intertidal reef flats at Ko Phuket, Thailand. Coral Reefs 1, 131–139.
Brown, B.E., Le Tissier, M.D.A., Scoffin, T.P. & Tudhope, A.W. 1990. Evaluation of the environmental impact
of dredging on intertidal coral reefs at Ko Phuket, Thailand, using ecological and physiological
parameters. Marine Ecology Progress Series 65, 273–281.
Brown, B.E. & Phongsuwan, N. 2004. Constancy and change on shallow reefs around Laem Pan Wa, Phuket,
Thailand over a 20-year period. Phuket Marine Biological Center Research Bulletin 65, 61–73.
Buddemeier, R.W. & Kinzie, R.A. 1976. Coral growth. Oceanography and Marine Biology: An Annual Review
14, 183–225.
Burnett, W.J. 2002. Longitudinal variation in algal symbionts (zooxanthellae) from the Indian Ocean zoanthid
Palythoa caesia. Marine Ecology Progress Series 234, 105–109.
Chansang, H., Phongsuwan, N. & Boonyanate, P. 1992. Growth of corals under effect of sedimentation along
the northwest coast of Phuket Island, Thailand. Proceedings of the Seventh International Coral Reef
Symposium, Guam 1, 241–248.
Chansang, H., Satapoomin, U. & Poovachiranon, S. (eds) 1999. Maps of Coral Reefs in Thai waters: Volume 2,
Andaman Sea. Bangkok: Coral Reef Resource Management Project, Department of Fisheries.
Clarke, K.R., Warwick, R.M. & Brown, B.E. 1993. An index showing breakdown of seriation, related to
disturbance, in a coral-reef assemblage. Marine Ecology Progress Series 102, 153–160.
Curray, J.R. 2005. Tectonics and history of the Andaman Sea region. Journal of Asian Earth Sciences 25,
187–232.
Darwin, C. 1842. The Structure and Distribution of Coral Reefs. London: Smith, Elder and Co.
Department of Marine and Coastal Resources, Thailand. 2005. Report on the Effects of the Indian Ocean
Tsunami in Thai Waters [in Thai]. Bangkok: Department of Marine and Coastal Resources.
Desai, B.N., Bhargava, R.M.S., Sarupia, J.S. & Pankajakshan, T. 1988. Oceanographic coverage of the
Andaman and Nicobar Seas. Indian Association for Advancement of Science, Island Development
Technology Options, 86–102.
Downs, C.A., Mueller, E., Phillips, S., Fauth, J.E. & Woodley, C.M. 2000. A molecular biomarker system for
assessing the health of a coral (Montastraea faveolata) during heat stress. Marine Biotechnology 2,
533–544.
Drew, E.A. 1972. The biology and physiology of algal invertebrate symbiosis. II. The density of algal cells
in a number of hermatypic hard corals and alcyonarians from various depths. Journal of Experimental
Dunne, R.P. & Brown, B.E. 2001. The influence of solar radiation on bleaching of shallow water reef corals
in the Andaman Sea, 1993–1998. Coral Reefs 20, 201–210.
Goodson, M.S. 2000. Symbiotic algae: molecular diversity in marginal coral reef habitat. Ph.D. thesis,
University of York, U.K.
Guzman-Speziale, M. & Ni, J.F. 1993 The opening of the Andaman Sea: where is the short term displacement
being taken up? Geophysical Research Letters 20, 2949–2952.
191
BARBARA E. BROWN
Hall, R. 1998. The plate tectonics of Cenozoic SE Asia and the distribution of land and sea. In Biogeography
and Evolution of SE Asia, R. Hall & J.D. Holloway (eds). Leiden, The Netherlands: Backhuys
Publishers, 99–131.
Haneburth, T., Stattegger, K. & Grootes, P.M. 2000. Rapid flooding of the Sunda Shelf: a late-glacial sea-
level record. Science 288, 1033–1035.
Hesp, P.A., Hung, C.C., Hilton, M., Loke Ming,C. & Turner, I.M. 1998. A first tentative Holocene sea level
curve for Singapore. Journal of Coastal Research, 14, 308–314.
Hoegh Guldberg, O. 1999 Climate change: coral bleaching and the future of the world’s coral reefs. Marine
and Freshwater Research 50, 839–866.
Hyder, P., Jeans, D.R.G., Cauquil, E.& Nerzic, R. 2005. Observations and predictability of internal solitons
in the northern Andaman Sea. Applied Ocean Research, 27, 1–11.
IPCC. 2001. Climate Change 2001 Synthesis report. R.T. Watson and Core Writing Team (eds). Geneva,
Switzerland: Intergovernmental Panel on Climate Change.
Janekarn, V. & Hylleberg, J. 1989. Coastal and offshore production along the west coast of Thailand (Andaman
Sea) with notes on physical-chemical variables. Phuket Marine Biological Center Research Bulletin
51, 1–20.
Johnstone, I. 1980. The ultrastructure of skeletogenesis in hermatypic corals. International Reviews of Cytology
67, 171–214.
Khan, P.K. & Chakraborty, P.P. 2005. Two-phase opening of Andaman Sea: a new seismotectonic insight.
Earth and Planetary Science Letters 229, 259–271.
Khokiattiwong, S. 1991. Coastal hydrography of the southern Andaman Sea of Thailand, Trang to Satun
province. In Proceedings of the First Workshop of the Tropical Marine Mollusc Programme (TMMP).
Phuket: Phuket Marine Biological Center, 87–93.
LaJeunesse, T.C. 2001. Investigating the biodiversity, ecology and phylogeny of endosymbiotic dinoflagellates
in the genus Symbiodinium using the ITS region: in search of a ‘species’ level marker. Journal of
LaJeunesse, T.C., Loh, W.K.W., van Woesik, R., Hoegh-Guldberg, O., Schmidt, G.W. & Fitt, W.K. 2003. Low
symbiont diversity in southern Great Barrier Reef corals, relative to those of the Caribbean. Limnology
Lough, J.M. & Barnes, D.J. 2000. Environmental controls on growth of the massive coral Porites. Journal of
Madhaven, B.B., Venkataraman, G., Shah, D. & Mohan, B.K. 1997. Revealing geology of the Great Nicobar
Island, Indian Ocean, by interpretation of airborne synthetic aperture radar images. International
Journal of Remote Sensing 18, 2723–2742.
McManus, J.W. 1985. Marine speciation, tectonics and sea-level changes in Southeast Asia. Proceedings of
the Fifth International Coral Reef Congress, Tahiti 4, 133–138.
Myers, R.F. 1991. Micronesian Reef Fishes: A Practical Guide to the Identification of Inshore Fishes of the
Tropical and Western Pacific. Guam: Coral Graphics, 2nd edition.
Nielsen, T.G., Bjornsen, P.K., Boonruang, P., Fryd, M., Hansen, P.J., Janekarn, V., Limtrakulvong, V., Munk,
P., Hansen, O.S., Satapoomin, S., Sawangarreruks, S., Thomsen, H.A. & Ostergaard, J.B. 2004.
Hydrography, bacteria and protist communities across the continental shelf and shelf slope of the
Andaman Sea (NE Indian Ocean). Marine Ecology Progress Series 274, 69–86.
Oldham, R.D. 1884. Note on the earthquake of 31 December 1881. Records of the Geological Survey of India
17, 45–53.
Osborne, A.R. 1990. The generation and propagation of internal solitons in the Andaman Sea. In Soliton
Theory: A Survey of Results, A.P. Fordy (ed.). Manchester: Manchester University Press, 152–173.
Osborne, A.R. & Burch, T.L. 1980. Internal solitons in the Andaman Sea. Science 208, 451–460.
Pal, T., Chakraborty, P.P., Gupta, T.D. & Singh, C.D. 2003. Geodynamic evolution of the outer-arc-forearc
belt in the Andaman Islands, the central part of the Burma-Java subduction complex. Geological
Magazine 140, 289–307.
Panutrakul, S. 1996. Water quality in Phuket Bay. Phuket Marine Biological Research Center Research Bulletin
61, 67–81.
192
Phongsuwan, N. 1991. Recolonization of a coral reef damaged by a storm on Phuket Island, Thailand. Phuket
Marine Biological Center Research Bulletin 56, 75–83.
Phongsuwan, N. Geographic Maps of Natural Resources in Southern Thailand [in Thai]. Bangkok: Department
of Marine and Coastal Resources (in press).
Phongsuwan, N. & Chansang, H. 1992. Assessment of coral communities in the Andaman Sea (Thailand).
Proceedings of the Seventh International Coral Reef Symposium 1, 114–121.
Pillai, C.S.G. 1972. Stony corals of the seas around India. Proceedings of a Symposium on Corals and Coral
Reefs. Marine Biological Association of India, 191–216.
Pillai, C.S.G. 1983. Coral reefs and their environs. In Mariculture Potential of Andaman and Nicobar Islands —
An Indicative Survey. Central Marine Fisheries Research Institute Bulletin, Cochin, India, 34, 36–43.
Potemra, J.T., Luther, M.E. & O’Brien, J.J. 1991. The seasonal circulation of the upper ocean in the Bay of
Bengal. Journal of Geophysical Research 96, 12,667–12,683.
Potts, D.C. 1983. Evolutionary disequilibrium among Indo-Pacific corals. Bulletin of Marine Science 33, 619–632.
Potts, D.C. & Jacobs, J.R. 2000. Evolution of reef-building scleractinian corals in turbid environments: a
paleo-ecological hypothesis. Proceedings of the Ninth International Coral Reef Symposium, Bali 1,
249–254.
Pubellier, M., Ego, F., Chamot-Rooke, N. & Rangin, C. 2003. The building of pericratonic mountain ranges:
structural and kinematic constraints applied to GIS-based reconstruction of SE Asia. Bulletin de la
Societé Geologique de France 174, 561–584.
Ramaswamy, V., Rao, P.S., Rao, K.H., Thwin, S., Rao, N.S. & Raiker, V. 2004. Tidal influence on suspended
sediment distribution and dispersal in the northern Andaman Sea and Gulf of Martaban. Marine
Geology 208, 33–42.
Rao, T.S.S. & Griffiths, R.C. 1998. Understanding the Indian Ocean. Perspectives on Oceanography Series.
Paris: UNESCO Publishing.
Reddiah, K. 1977. The coral reefs of the Andaman and Nicobar Islands. Records of the Zoological Survey of
India 72, 315–324.
Rodolfo, K.S. 1969. Bathymetry and marine geology of the Andaman Basin and tectonic implications for
Southeast Asia. Geological Society of America Bulletin 80, 1203–1230.
Rosen, B.R. 1975. The distribution of reef corals. Report of the Underwater Association 1, 1–16.
Rowan, R. 2004. Thermal adaptation in reef coral symbionts. Nature 430, 742.
Satapoomin, U. 1993. Responses of corals and coral reef to the 1991 coral reef bleaching event in the Andaman
Sea, Thailand. M.Sc. thesis, Chulalongkorn University, Thailand.
Satapoomin, U. 2002. Comparative Study of Reef Fish Fauna in Thai Waters: The Gulf of Thailand versus
the Andaman Sea. Technical Paper Phuket Marine Biological Center.
Satapoomin, U., Phongsuwan, N. & Brown, B.E. 2006. A preliminary synopsis of the effects of the Indian
Ocean tsunami on the coral reefs of western Thailand. Phuket Marine Biological Center Research
Bulletin 67, 77–80.
Scheer, G. 1971. Coral reefs and coral genera in the Red Sea and Indian Ocean. Symposium of the Zoological
Society of London 28, 329–367.
Scoffin, T.P. & Le Tissier, M.D.A. 1998. Late Holocene sea level and reef-flat progradation, Phuket, South
Thailand. Coral Reefs 17, 273–276.
Scoffin, T.P., Tudhope, A.W., Brown, B.E., Chansang, H. & Cheeney, R.F. (1992) Patterns and possible
environmental controls of skeletogenesis of Porites lutea, South Thailand. Coral Reefs 11, 1–11.
Searle, M. 2006. Co-seismic uplift of coral reefs along the western Andaman Islands during the December
26, 2004 earthquake. Coral Reefs, 25, 2.
Sewell, R.B.S. 1925. The geography of the Andaman Sea basin. Memoirs of the Asiatic Society of Bengal 9,
1–26.
Sewell, R.B.S. 1935. Studies on coral and coral formations in Indian waters. Memoirs of the Asiatic Society
of Bengal 9, 461–540.
Sieh, K. 2005. What happened and what’s next? Nature 434, 573–574.
Soegiarto, A. 1985. Oceanographic assessment of the East Asian Seas. Environment and Resources in the
Pacific, UNEP Regional Seas Reports and Studies 69, 173–184.
193
BARBARA E. BROWN
Soegiarto, A. & Birowo, A.T. 1975. Oceanologic Atlas of the Indonesian and Adjacent Waters. Volume 1.
Jakarta, Indonesia: National Institute of Oceanology.
Spalding, M.D., Ravilious, C. & Green, E.P. 2001. World Atlas of Coral Reefs. Berkeley: University of
California Press.
Spencer, T. Coral reefs and the tsunami of 26 December 2004: generating processes and ocean-wide patterns
of impact. Atoll Research Bulletin (in press).
Stambler, N., Popper, N., Dubinsky, Z. & Stimson, J. 1991. Effects of nutrient enrichment and water motion
on the coral Pocillopora damicornis. Pacific Science 45, 299–307.
Stanley, G.D. 1988. The history of early Mesozoic reef communities: a three step process. Palaios 3, 170–183.
Suharsono & Soekarno. 1983. Kandungan zooxanthella pada karang batu di terumbu karang Pulau Pari.
Oseanologi Indonesia 16, 1–7.
Tija, H.D. 1980. The Sunda Shelf, Southeast Asia. Zeitchrift fur Geomorphologie 24, 405–427.
Tomczak, M. & Godfrey, J.S. 1994. Regional Oceanography: An Introduction. Oxford, U.K.: Pergamon.
Tudhope, A.W. & Scoffin, T.P. 1994. Growth and structure of fringing reefs in a muddy environment, South
Thailand. Journal of Sedimentary Research A64, 752–764.
Turak, E., Veron, C. & Sanpanich, K. 2005. Coral reef status after 26 December 2004 tsunami. In Final
Technical Report Rapid Assessment Survey of Tsunami-Affected Reefs of Thailand, R.A. Allen & G.S.
Stone (eds). Boston: New England Aquarium, 16–30.
Turner, J.R., Vousden, D., Klaus, R., Satyanarayan, C., Fenner, D., Venkataraman, K., Rajan, P.T. & Subba
Rao, N.V. 2001. Report of Phase 1: Remote Sensing and Rapid Site Assessment Survey April 2001.
Coral Reef Systems of the Andaman Islands, Government of India and United Nations Development
Programme, Global Environment Facility, New Delhi, India.
UNEP/IUCN. 1988. Coral Reefs of the World Volume 2: Indian Ocean, Red Sea and Gulf. UNEP Regional
Seas Directories and Bibliographies. Gland, Switzerland: IUCN and Nairobi, Kenya: UNEP.
Varkey, M.J. 1986. Salt balance and mixing in the Bay of Bengal. Ph.D. thesis, University of Kerala, India.
Varkey, M.J., Murty, V.S.N. & Suryanarayana, A. 1996. Physical oceanography of the Bay of Bengal and
Andaman Sea. Oceanography and Marine Biology: An Annual Review 34, 1–70.
Wallace, C.C. 1999. Staghorn Corals of the World: A Revision of the Coral Genus Acropora. Collingwood,
Australia: CSIRO Publishing.
Wallace, C.C. & Muir, P.R. 2005. Biodiversity of the Indian Ocean from the perspective of staghorn corals
(Acropora spp.). Indian Journal of Marine Science 31, 42–49.
Webster, P.J, Moore, A.M., Loschnigg, J.P. & Lebel, R.R. 1999. Coupled ocean-atmosphere dynamics in the
Indian Ocean during 1997–1998. Nature 401, 356–360.
Wilkinson, C. 2004. Status of Coral Reefs of the World 2004. Townsville: Australian Institute of Marine Science.
Wilson, M.E.J. & Rosen, B.R. 1998. Implications of paucity of corals in the Paeogene of SE Asia: plate
tectonics or centre of origin? In Biogeography and Geological Evolution of SE Asia, R. Hall & J.D.
Holloway (eds). Leiden, The Netherlands: Backhuys Publishers, 165–195.
Wyrtki, K. 1973. An equatorial jet in the Indian Ocean. Science 181, 262–264.
Yesaki, M. & Jantarapagdee, P. 1981. Wind stress and sea temperature changes off the west coast of Thailand.
In Special Publication on the Occasion of the 10th Anniversary of Phuket Marine Biological Centre.
Phuket, Thailand: Phuket Marine Biological Center, 27–42.
194
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Taylor & Francis
THE HUMBOLDT CURRENT SYSTEM

OF NORTHERN AND CENTRAL CHILE
OCEANOGRAPHIC PROCESSES, ECOLOGICAL INTERACTIONS
AND SOCIOECONOMIC FEEDBACK
MARTIN THIEL1,2, ERASMO C. MACAYA1, ENZO ACUÑA1, WOLF E. ARNTZ3, HORACIO

BASTIAS1, KATHERINA BROKORDT1,2, PATRICIO A. CAMUS4,5, JUAN CARLOS
CASTILLA5,6, LEONARDO R. CASTRO7, MARITZA CORTÉS1, CLEMENT P. DUMONT1,2,
RUBEN ESCRIBANO8, MIRIAM FERNANDEZ5,9, JHON A. GAJARDO1, CARLOS F.
GAYMER1,2, IVAN GOMEZ10, ANDRÉS E. GONZÁLEZ1, HUMBERTO E. GONZÁLEZ10,
PILAR A. HAYE1,2, JUAN-ENRIQUE ILLANES1, JOSE LUIS IRIARTE11, DOMINGO A.
LANCELLOTTI1, GUILLERMO LUNA-JORQUERA1,2, CAROLINA LUXORO1,
PATRICIO H. MANRIQUEZ10, VÍCTOR MARÍN12, PRAXEDES MUÑOZ1, SERGIO A.
NAVARRETE5,9, EDUARDO PEREZ1,2, ELIE POULIN12, JAVIER SELLANES1,8, HECTOR
HITO SEPÚLVEDA13, WOLFGANG STOTZ1, FADIA TALA1, ANDREW THOMAS14,
CRISTIAN A. VARGAS15, JULIO A. VASQUEZ1,2 & J.M. ALONSO VEGA1,2
1
Facultad de Ciencias del Mar, Universidad Católica del Norte,
Larrondo 1281, Coquimbo, Chile
E-mail: thiel@ucn.cl
2Centro de Estudios Avanzados en Zonas Áridas (CEAZA), Coquimbo, Chile
3Alfred Wegener Institute for Polar and Marine Research, Columbusstrasse,
27568 Bremerhaven, Germany

4Facultad de Ciencias, Universidad Católica de la Santísima Concepción,
Casilla 297, Concepción, Chile

5Center for Advanced Studies in Ecology and Biodiversity (CASEB), Santiago, Chile
6Facultad de Ciencias Biológicas, Pontificia Universidad Católica de Chile,
Casilla 114-D, Santiago, Chile

7Laboratorio de Oceanografía Pesquera y Ecología Larval (LOPEL),
Departamento de Oceanografía, Universidad de Concepción, Concepción, Chile

8Center for Oceanographic Research in the Eastern South Pacific (COPAS),
Departamento de Oceanografía, Facultad de Recursos Naturales y Oceanografía,

Universidad de Concepción, Estación de Biología Marina, PO Box 42, Dichato, Chile
9Coastal Marine Research Station, Departamento de Ecología,
Pontificia Universidad Católica de Chile, Casilla 114D, Santiago, Chile

10Instituto de Biología Marina, Universidad Austral de Chile,
PO Box 567, Campus Isla Teja, Valdivia, Chile

11Instituto de Acuicultura, Universidad Austral de Chile, PO Box 1327, Puerto Montt, Chile
12Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile,
Las Palmeras 3425, Casilla 653, Nuñoa, Santiago, Chile

13Departamento de Geofísica, Facultad de Ciencias Físicas y Matemáticas,
Universidad de Concepción, Casilla 160-C, Concepción, Chile

14School of Marine Sciences, University of Maine, Orono, Maine 04469-5741, U.S.
15Unidad de Sistemas Acuáticos, Centro de Ciencias Ambientales EULA,
Universidad de Concepción, Concepción, Chile
195
MARTIN THIEL ET AL.
Abstract The Humboldt Current System (HCS) is one of the most productive marine ecosystems
on earth. It extends along the west coast of South America from southern Chile (~42°S) up to
Ecuador and the Galapagos Islands near the equator. The general oceanography of the HCS is
characterised by a predominant northward flow of surface waters of subantarctic origin and by
strong upwelling of cool nutrient-rich subsurface waters of equatorial origin. Along the coast of
northern and central Chile, upwelling is localised and its occurrence changes from being mostly
continuous (aseasonal) in northern Chile to a more seasonal pattern in southern-central Chile.
Several important upwelling centres along the Chilean coast are interspersed with long stretches
of coast without or with sporadic and less intense upwelling. Large-scale climatic phenomena
(El Niño Southern Oscillation, ENSO) are superimposed onto this regional pattern, which results
in a high spatiotemporal heterogeneity, complicating the prediction of ecological processes along
the Chilean coast. This limited predictability becomes particularly critical in light of increasing
human activities during the past decades, at present mainly in the form of exploitation of renewable
resources (fish, invertebrates and macroalgae). This review examines current knowledge of ecolog-
ical processes in the HCS of northern and central Chile, with a particular focus on oceanographic
factors and the influence of human activities, and further suggests conservation strategies for this
high-priority large marine ecosystem. Along the Chilean coast, the injection of nutrients into surface
waters through upwelling events results in extremely high primary production. This fuels zoo-
plankton and fish production over extensive areas, which also supports higher trophic levels,
including large populations of seabirds and marine mammals. Pelagic fisheries, typically concen-
trated near main upwelling centres (20–22°S, 32–34°S, 36–38°S), take an important share of the
fish production, thereby affecting trophic interactions in the HCS. Interestingly, El Niño (EN) events
in northern Chile do not appear to cause a dramatic decline in primary or zooplankton production
but rather a shift in species composition, which affects trophic efficiency of and interactions among
higher-level consumers. The low oxygen concentrations in subsurface waters of the HCS (oxygen-
minimum zone, OMZ) influence predator-prey interactions in the plankton by preventing some
species from migrating to deeper waters. The OMZ also has a strong effect on the bathymetric
distribution of sublittoral soft-bottom communities along the Chilean coast. The few long-term
studies available from sublittoral soft-bottom communities in northern and central Chile suggest
that temporal dynamics in abundance and community composition are driven by interannual phe-
nomena (EN and the extent and intensity of the OMZ) rather than by intra-annual (seasonal) patterns.
Macrobenthic communities within the OMZ are often dominated in biomass by sulphide-oxidising,
mat-forming bacteria. Though the contribution of these microbial communities to the total primary
production of the system and their function in structuring OMZ communities is still scarcely known,
they presumably play a key role, also in sustaining large populations of economically valuable
crustaceans. Sublittoral hard bottoms in shallow waters are dominated by macroalgae and suspen-
sion-feeder reefs, which concentrate planktonic resources (nutrients and suspended matter) and
channel them into benthic food webs. These communities persist for many years and local extinc-
tions appear to be mainly driven by large-scale events such as EN, which causes direct mortality
of benthic organisms due to lack of nutrients/food, high water temperatures, or burial under
terrigenous sediments from river runoff. Historic extinctions in combination with local conditions
(e.g., vicinity to upwelling centres or substratum availability) produce a heterogeneous distribution
pattern of benthic communities, which is also reflected in the diffuse biogeographic limits along
the coast of northern-central Chile. Studies of population connectivity suggest that species with
highly mobile planktonic dispersal stages maintain relatively continuous populations throughout
most of the HCS, while populations of species with limited planktonic dispersal appear to feature
high genetic structure over small spatial scales. The population dynamics of most species in the
HCS are further influenced by geographic variation in propagule production (apparently caused by
local differences in primary production), by temporal variation in recruit supply (caused by upwelling
196
THE HUMBOLDT CURRENT SYSTEM OF NORTHERN AND CENTRAL CHILE
events, frontal systems and eddies), and topographically driven propagule retention (behind head-
lands, in bay systems and upwelling shadows). Adults as well as larval stages show a wide range
of different physiological, ecological and reproductive adaptations. This diversity in life-history
strategies in combination with the high variability in environmental conditions (currents, food
availability, predation risk, environmental stress) causes strong fluctuations in stocks of both
planktonic and benthic resources. At present, it remains difficult to predict many of these fluctua-
tions, which poses particular challenges for the management of exploited resources and the con-
servation of biodiversity in the HCS. The high spatiotemporal variability in factors affecting
ecological processes and the often-unpredictable outcome call for fine-scale monitoring of recruit-
ment and stock dynamics. In order to translate this ecological information into sustainable use of
resources, adaptive and co-participative management plans are recommended. Identification of areas
with high biodiversity, source and sink regions for propagules and connectivity among local
populations together with developing a systematic conservation planning, which incorporates deci-
sion support systems, are important tasks that need to be resolved in order to create an efficient
network of Marine Protected Areas along the coast of northern-central Chile. Farther offshore, the
continental shelf and the deep-sea trenches off the Chilean coast play an important role in bio-
geochemical cycles, which may be highly sensitive to climatic change. Research in this area should
be intensified, for which modern research vessels are required. Biodiversity inventories must be
accompanied by efforts to foster taxonomic expertise and museum collections (which should
integrate morphological and molecular information). Conservation goals set for the next decade
can only be achieved with the incorporation of local stakeholders and the establishment of efficient
administrative structures. The dynamic system of the HCS in northern-central Chile can only be
understood and managed efficiently if a fluent communication between stakeholders, administrators,
scientists and politicians is guaranteed.
Introduction
The deep-blue colour of the water observed for a long time past gave place to a green colour, and on
the whole there was a great change in the general character of the surface fauna, pointing to the nearness
of a great continent, similar to what was observed off Japan and elsewhere. On November 18 [1875],
the water was very green in colour, and the ship occasionally passed through large red or brown patches,
which the tow-net showed to be due to immense numbers of red copepods, hyperids, and other Crustacea.
Murray (1895), on approaching Valparaíso aboard H.M.S. Challenger
Such a vivid language was rarely used by John Murray to refer to the abundance of planktonic
organisms in surface waters. Only for the surface plankton from the Agulhas Bank off South Africa
did he employ similar colourful language, referring to “myriads of Zoeae and a few larger Mega-
lopae”. Scientists studying the plankton ecology of the Eastern Boundary Currents (EBCs) are used
to the sight of these dense accumulations of zooplankton, which are often found in sharply defined
patches. It is the intense upwelling of nutrient-rich waters in the EBCs that fuels the extraordinary
high primary production (PP) in the EBCs, which forms the basis of the food web supporting some
of the largest fisheries of the world. However, the frequency and intensity of upwelling within the
EBCs varies, mainly depending on large-scale climatic forcing, latitudinal/seasonal signals and
local factors, such as the width of the shelf, coastal topography, and sources of upwelled waters
(Thomas et al. 2004). Although the overall importance of upwelling in these large marine ecosys-
tems is relatively well known, the effects of temporal and spatial variability of upwelling on the
ecology and productivity of the planktonic and benthic communities remain poorly understood.
Herein these effects are explored, using the Humboldt Current System (HCS), one of the most
197
MARTIN THIEL ET AL.
productive EBCs (Halpin et al. 2004, Montecino et al. 2005), as a model system. In the present
paper knowledge on the HCS in northern-central Chile is reviewed because this area is characterised
by a complex set of temporally changing and geographically variable conditions that represent a
particular challenge to science and management (see also Artisanal benthic fisheries and following
sections, p. 278ff). The aims of this review are 4-fold: to (1) review current knowledge, (2) reveal
gaps in information, (3) indicate conservation priorities, and (4) propose future research avenues.
The HCS, by some authors named the Peru-Chile Current System, extends from ~42°S up to
about the equator (Montecino et al. 2005). The main oceanographic features of this system are
often described as cold nutrient-rich waters being transported northward and nutrient-enriched
subsurface waters upwelled along the shorelines of Ecuador, Peru and northern Chile. Occasionally
the nutrient-supply engine of the HCS is interrupted by influx of warm and nutrient-depleted
equatorial waters; during such events (El Niño, EN) the northward flow of cool nutrient-rich waters
is suppressed and upwelling intensity is often reduced (W. Palma et al. 2006). Individual cycles of
this El Niño Southern Oscillation (ENSO) last for several years, but predictability of EN events is
still very limited. On shorter temporal scales, there is a seasonal (predictable) pattern of climate
and oceanography at high latitudes, which becomes aseasonal (and less predictable) at mid- and
low latitudes (Blanco et al. 2001, Carr et al. 2002). It can be argued that the predictability of
oceanographic conditions is lowest in the mid-region of the HCS (from 18°S to about 32°S; see
also Thomas et al. 2001a) because here the seasonal variations are occasionally overshadowed by
the interannual ENSO cycles, whereas at low latitudes (<18°S) the conditions may remain relatively
stable for several years in a row, only being disturbed, but then severely, by EN events. To add to
this variability, the HCS between 18°S and ~40°S features a complex coastal oceanography, where
the main equatorward current is enveloped by a set of counter- and undercurrents, the width, location
and intensity of which also vary in time. Herein we focus on this area between 18°S and ~40°S
because the interaction between seasonal and interannual (ENSO) signals results in high temporal
variability, which affects the ecological processes in this region.
Overall, the coastline of northern and central Chile (18° to ~40°S) is relatively straight
(Figure 1), but in the nearshore region small-scale geographic features produce a high spatial
heterogeneity, which also influences oceanographic conditions in this area. Several bay systems
are found along the coast of northern-central Chile. Circulation in these bays is complex with
counterrotating gyres (Valle-Levinson et al. 2000) affecting larval transport and settlement patterns
(A.T. Palma et al. 2006), most likely related to depth- and site-dependent retention or export
scenarios (Yannicelli et al. 2006a). Headlands favour the generation of powerful coastal flow
structures (squirts) transporting surface waters up to 100 km offshore (Marín et al. 2003a). In
contrast, long stretches of exposed outer coast without headlands or bays, as found for example in
northern Chile between 20° and 22°S, favour alongshore currents leading to relatively homogeneous
conditions and downstream transport (W. Palma et al. 2006).
Numerous studies have examined the effects of oceanographic conditions on a variety of
ecological processes in the HCS (e.g., Carrasco & Santander 1987, Tarazona et al. 1988a,b, Alheit &
Niquen 2004, Arntz et al. 2006). Upwelling or ENSO-related conditions affect PP (Wieters et al.
2003, Iriarte & González 2004), zooplankton community composition (Escribano et al. 2004a), fish
population dynamics (Halpin et al. 2004), dispersal of larvae (Poulin et al. 2002a,b), growth of
benthic algae (Wieters 2005), benthic-pelagic coupling (Graco et al. 2006), population dynamics
of benthos organisms (Castilla & Camus 1992), and a variety of other processes. Most of these
studies have focused on direct cause-effect relationships between oceanographic factors and eco-
logical responses. Recent studies in central Chile indicate complex interactions between upwelling,
supply of recruits, and grazers or predators (Nielsen & Navarrete 2004, Wieters 2005). These studies
show the importance of bottom-up and top-down processes, which may vary on small temporal
and spatial scales, making predictions difficult. In this context, Navarrete et al. (2002) emphasised
198
Fishery unit CHILE

18° Arica
20° Iquique/Rio Loa Iquique

I - II
Northern
Mejillones
Antofagasta Antofagasta
25°
HU
Caldera
III - IV
M
Isla Choros
BO
30° Coquimbo Coquimbo
D
L
(Lengua de Vaca) T
Punta Curaumilla Valparaíso
CU
Central
central Chile
RREN T SYSTEM
35°
V - IX Concepcin Concepción
Golfo Arauco
Valdivia
40°
Chiloé
Southern
45°
X - XII
50°
55°
Figure 1 Study region in northern Chile with the main upwelling regions and localities mentioned throughout
the text. Principal upwelling centers in black dots, other sites with frequent upwelling indicated with grey
dots, coastal stretches with occasional upwelling shown as thick black line.
that “lack of consistent trends among sites … shows that El Niño effects on interannual recruitment
variation are not predictable”. Not only marine biologists but above all the organisms inhabiting the
HCS are grappling with this limited predictability. How do these organisms deal with the difficulty
of foreseeing the availability of dispersal windows, food, competitors or predators in the near
future? Many of the following sections will explicitly or implicitly address this question. The main
focus, though, will be on the outcomes of ecological processes in the HCS, the processes that
govern them and their relevance in a socioeconomic and conservation context.
Variable distribution patterns and species interactions in the HCS are not only due to oceano-
graphic processes but are also increasingly affected by human activities that reduce the abundance
of some species while favouring others. When humans first started to use the natural resources of
the HCS, they opportunistically reacted to the system and exploited natural resources where these
were available and accessible. When a particular resource became scarce or inaccessible they either
searched for the resource in other places or shifted to alternative resources (e.g., Llagostera 1979,
Méndez & Jackson 2004). With increasing population pressure and technological advances, human
pressure on the HCS has intensified. Wave-sheltered bays along the coast with dense human popu-
lations are impacted by intense shipping traffic, artificial coastline constructions and wastewater
influx (Fernández et al. 2000). Mining and agriculture activities have resulted in severe contamination
of some coastal areas, fishing pressure has intensified and extended into previously inaccessible
regions and zones and finally climate change (increased ultraviolet (UV) radiation, global warming)
has also reached the HCS. From being opportunistic users who responded to natural variations in
resource abundance, humans have now become important actors who directly affect many of the
199
MARTIN THIEL ET AL.
natural processes in the HCS. Recognition of this impact has called increasingly for planning and
regulation of human activities. The limited predictability of oceanographic conditions and the
ecological processes in the HCS, however, represents an enormous challenge for efficient manage-
ment of this large marine ecosystem. Herein we strive to describe those aspects of the system that
are comparatively predictable and to identify those that will require additional knowledge before
they can be reliably predicted. In order to achieve this goal, this review provides (1) a brief overview
of historic research activities, (2) a description of the main oceanographic conditions in the HCS of
northern and central Chile, (3) an analysis of the pelagic environment and top consumers, (4) an
introduction to benthic systems and nearshore biogeography, (5) a discussion of biological adap-
tations of organisms, (6) a portrayal of socioeconomic aspects related to the exploitation of natural
resources, and finally (7) a scenario for marine conservation and an outlook identifying some of
the main administrative and scientific tasks for the future.
A brief history of research on the Humboldt current system

First reports on a cool current in the southeastern Pacific
First written accounts of a cold current along the western coast of South America come from the
European explorers. According to Gunther (1936), the history of the Humboldt Current began in
1515 when Vasco Nuñez de Balboa first sighted the South Sea. The knowledge of this oceanic
current dates to the sixteenth century when several Europeans carried out observations suggesting
that waters were cool: Pascual de Andagoya in 1522 was the first Spaniard to explore to the south
of the Panamanian coast, Agustin de Zarate published in 1555 his Historia del descubrimiento y
Conquista de la Provincia del Peru, and the Jesuit José de Acosta described in his Historia Natural y
Moral de Indias (1591) the temperature conditions and their influence on the climate (Santibáñez
1944). Other travellers carried out additional observations (for details see Gunther 1936), but it
was the German naturalist Alexander von Humboldt who took the first temperature measurements
(Gunther 1936, Santibáñez 1944). Humboldt (1846), in his book Cosmos, wrote on page 301: “in
the Southern Pacific Ocean, … a current the effect of whose low temperature on the climate of the
adjacent coast was first brought into notice by myself in the autumn of 1802. This current brings
the cold water of high southern latitudes to the coast of Chile, and follows its shores and those of
Peru northward”. Additionally, Humboldt also was the first to describe EN temperature anomalies
in coastal waters (Kortum 2002).
From natural history to ecology to marine conservation

The first Chilean naturalist was Abate Juan Ignacio Molina. He published the following works in
Italy: Compendio della storia geografica, naturale, e civili del regno del Chile (1776) and Saggio
sulla storia naturale del Chili (1782), which include detailed descriptions of the mineral wealth
and native flora and fauna based on the Linnaean system of classification (Ronan 2002). Claudio
Gay, a French naturalist who arrived in Chile in 1828, as a result of 12 yr of exploring Chile under
government contract, wrote his Historia fisica y política de Chile. In 29 volumes Gay described
the history, botany and zoology of the country, including detailed descriptions of the marine fauna
(e.g., 79 species of decapod crustaceans) (Jara 1997). Rodulfo Amando Philippi, who emigrated
from Germany to Chile in 1851, was a physician by profession but a naturalist by inclination, and
he is considered one of the most recognised and influential scientists in the development of natural
sciences in Chile (Castro et al. 2006). He contributed numerous taxonomic descriptions to the
knowledge of biological diversity of Chile. In a recent article, Castro et al. (2006) conclude that
in comparison with other taxonomists, Philippi was the author of the largest number of presently
200
valid species in the Chilean biota. Other scientists who contributed enormously to the knowledge
of the fauna and flora from the HCS of northern-central Chile included Alcide D’Orbigny, Eduard
Friedrich Poeppig and Carlos Emilio Porter Mosso.
Most expeditions during the eighteenth and nineteenth centuries produced observations and
collections that were gathered by individual scientists or ‘naturalists’ aboard global voyages of
exploration and discovery. These voyages often had political, military and economic purposes, with
science being a secondary or even incidental activity (Fiedler & Lavín 2006). After Humboldt’s
travels, several pioneer expeditions were carried out in the southern Pacific, but most of them
rounded Cape Horn on their way into the Pacific, stopped in Concepción or more commonly in
Valparaíso, and then turned off to Juan Fernández and from there progressed further into the
southwest Pacific. The coasts of northern Chile were rarely visited by these expeditions. Also the
U.S. Exploring Expedition (1838–1842), the main objective of which was to facilitate American
commerce (Johnson 1995), left the coast of northern Chile untouched on its way from Valparaíso
to Callao. Other expeditions took the same route on their return trips. For example, the Novara
Expedition passed through Valparaíso in April 1859, coming from Tahiti. The corvette H.M.S.
Challenger reached Valparaíso in November 1875 coming from Juan Fernández and then continued
to the south. Later expeditions, such as the Fisheries Commission Steamer Albatross, with Alexander
Agassiz on board, explored the northern parts of the HCS, but rarely reached farther south than
Callao, Peru. A notable exception to this general pattern was the H.M.S. Beagle (1831–1836), with
Charles Darwin aboard, which after leaving Valparaíso made stopovers in Coquimbo (30°S), Caldera
(27°S) and Iquique (20°S) before continuing directly to Callao (Peru) (Darwin 1851, 1854).
Expeditions conducted during the first half of the twentieth century were mainly dedicated to
the study of the local biodiversity. One of the most important expeditions during that time, the
Lund University Chile Expedition in 1948–1949, explored the Chilean coast between Iquique and
the Magellan Straits, but of the 277 stations visited, only 79 were located in northern and central
Chile (Brattström & Dahl 1951). Only during the last half of the twentieth century have concentrated
research efforts been directed toward the oceanography and ecology of the HCS in northern Chile
(e.g., Gallardo 1963 and many others).
While those initial studies have provided important information on the description of species
distribution and abundance in the HCS along the coast of Chile, research during the last quarter
of the twentieth century became much more process oriented (Castilla & Largier 2002, Escribano
et al. 2004a,b, Montecino et al. 2005 and citations therein). In particular during the past decade,
marine conservation has become an important topic in the marine sciences literature of Chile (see,
e.g., Castilla 1996, 2000, Fernández et al. 2000, Fernández & Castilla 2000, 2005, Moreno 2001).
Thus, as in other regions of the HCS (e.g., Pauly et al. 1989), research in the HCS along the Chilean
coast has shifted from a description of taxonomy and patterns to the examination of processes,
which has also resulted in an increasing trend of interdisciplinary studies, in particular between
ecologists and oceanographers.
Oceanographic conditions in the southeastern Pacific

The HCS is the equatorward-flowing, eastern portion of the basin-scale southeast Pacific anti-
cyclonic gyre, bounded to the north by the equatorial current system and to the south by the West
Wind Drift (WWD). The HCS is one of the four major global EBCs, characterised by dominant
equatorward alongshore wind stress, offshore Ekman transport, coastal upwelling of cold, nutrient-
rich subsurface water and highly productive fisheries (Hill et al. 1998). The narrow continental
shelf and meridionally oriented coastline of South America allow efficient transmittal of atmospheric
and oceanographic signals imposed from both lower latitudes through the equatorial current system
and from higher latitudes where increasing seasonality in both wind forcing and oceanic response
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is seen with increasing latitude. The coast can be divided into general zones based on physical
characteristics. The shelf (200 m) off Peru is up to 100 km wide. Along the Chilean coast, four
zones (Figueroa 2002) include the northern region (north of ~32°S) with little freshwater influence
and an extremely narrow shelf (<10 km), a widening shelf from 32°S to 36°S, a wider shelf (~70 km)
from 36°S to 42°S with increased freshwater influence, especially in winter, and a high-latitude
region (>42°S) with a wider topographically complex, fjord-indented coastline and with strong
freshwater influence.
Wind forcing in the southeast Pacific is dominated by the influence of the southeast Pacific
subtropical anticyclone that creates equatorward, upwelling-favourable winds along most of the
South American Pacific coast with latitudinally varying seasonality. Between 5°S and ~35°S,
monthly mean winds remain upwelling favourable throughout the year and are weakest along the
northern Chilean coast ~17–23°S (Shaffer et al. 1999, Thomas et al. 2001a). Distinct phase differ-
ences in the annual maxima are evident as a function of latitude (Thomas et al. 2001a), where
maximum upwelling strength shifts from austral autumn–winter off Peru (Bakun & Nelson 1991,
Carr et al. 2002), to austral spring–summer off northern Chile (Blanco et al. 2001) and austral
summer south of ~30°S (Shaffer et al. 1999). Upwelling is augmented by a decrease in alongshore
wind stress near shore that creates a region of cyclonic (negative) wind stress curl in the coastal
zone, lifting isotherms. Two regions of maximum alongshore wind stress are evident, centred at
~15°S and ~30°S (Shaffer et al. 1999, Thomas et al. 2001a, Hormazábal et al. 2004). North of
30°S, strong land–sea thermal contrast in austral summer enhances upwelling-favourable alongshore
wind stress (Rutllant et al. 2004a). At the very lowest latitudes (north of ~5°S), the annual meridional
migration of the Inter-Tropical Convergence Zone (ITCZ) creates both wind and precipitation (and
therefore stratification) seasonality. Beginning at ~35°S and extending poleward, seasonality in the
strength of the subtropical anticyclone creates seasonal reversals between an austral summer
upwelling maximum and mean winter conditions of poleward, downwelling-favourable winds
driven by winter storms associated with the polar front (Shaffer et al. 1999, Rutllant et al. 2004a).
The strength and duration of these winter conditions increases with latitude. South of ~45°S, events
associated with polar front activity create poleward alongshore monthly mean winds, downwelling
conditions and strong precipitation in coastal regions throughout the year.
The large-scale equatorward surface flow of the HCS within the basin-scale gyre belies a
complicated and still relatively poorly sampled flow structure closer to the coast. The following
description is a general overview of commonly observed characteristics reviewed in more detail
by Strub et al. (1998). The northern latitude at which the WWD approaches the continent shifts
seasonally from 35°S to 40°S in austral winter to ~45°S in austral summer. Flow from the WWD
branches north and south (Figure 2). The southern arm joins poleward coastal flow of the Cape
Horn Current, a buoyancy-driven coastal current with input from local river runoff, which is
strongest during austral winter. The northern arm forms the main flow of the Humboldt Current
that proceeds equatorward along the basin margin and joins the westward-flowing South Equatorial
Current at lowest latitudes, off northern Peru-Ecuador. This main flow of the Humboldt Current is
located seaward (~75–85°W) of a system of narrower coastal flows. Closest to the coast and coupled
to local wind forcing and coastal upwelling, the Chile Coastal Current (CCC) flows predominantly
equatorward. In summer, equatorward flow is most latitudinally extensive, traceable to 35–40°S
(Atkinson et al. 2002) and flowing along the entire central and northern Chilean coast, continuous
with the equatorward flowing Peru Coastal Current (PCC), also forced by local winds, as far as
~5°S. In winter, the PCC strengthens in association with the winter maximum in alongshore wind
stress. Off northern Chile (18–24°S) the seasonal maximum of the CCC is in the autumn (Blanco
et al. 2001), while south of ~25°S the CCC becomes increasingly seasonally variable. South of
~35°S, in association with the seasonal reversal in alongshore wind stress, the CCC is predominantly
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−92° −88° −84° −80° −76° −72° W
SEC 0°
PC
−10°
PC
C
CC
CCC −20°
P CCC
Southern latitude
HC
PCCC
−30°
CCC
−40°
WWD
−50°
CHC
Figure 2 Overview of the surface currents in the eastern South Pacific that influence the north-central Chilean
coast, showing the West Wind Drift (WWD), the main flow of the Humbolt Current (HC), the Cape Horn
Current (CHC), the Chile Coastal Current (CCC), the Peru Coastal Current (PCC), the Peru-Chile Counter-
current (PCCC) and the South Equatorial Current (SEC).
poleward in winter. Between these seasonal coastal currents and the more continuous equatorward
flow of the main Peru-Chile Current is the poleward-flowing Peru-Chile Countercurrent located
~100–300 km offshore, originating in equatorial regions likely as Equatorial Undercurrent water
and continues to ~35°S. This flow is poorly resolved in drifter data (Chaigneau & Pizarro, 2005)
but evident in altimeter data (Strub et al. 1995) and temperature-salinity characteristics (Strub et al.
1998). Beneath the coastal surface currents, a Poleward Undercurrent is continuous from northern
Peru to latitudes as high as 45–50°S (Silva & Neshyba 1979). This undercurrent, located over the
continental slope (Zuta & Guillén 1970, Shaffer et al. 1995, Leth et al. 2004), delivers relatively
saline, nutrient-rich, low-oxygen water (equatorial subsurface water, ESSW) to coastal regions; it
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is a primary contributor to upwelling off Peru and northern Chile (Huyer et al. 1987) and in Chilean
coastal regions at least as far as 35°S (Fonseca 1989).
From 5°S to ~40°S, the primary coastal oceanographic characteristics are isotherms and iso-
halines of the upper ~100 m tilted upward toward the coast and associated with upwelling. Off
Peru, the seasonally maximum winter winds and the small Coriolis term combine to create strong
offshore Ekman transport with upwelling effects evident at least as far as 400 km offshore. Along
the Chilean coast, four regions are recognised for especially strong upwelling, most likely due to
topographic enhancement by headlands, at Antofagasta (23°S), Coquimbo (30°S), Valparaíso (33°S)
and Concepción (37°S) (Figueroa & Moffat 2000, Mesías et al. 2003). Off southern-central Chile
(36–40°S), coastal stratification imposed by freshwater runoff becomes important even during
summer upwelling conditions (Atkinson et al. 2002).
Strong interannual variability is superimposed on mean seasonal patterns by ENSO signals
propagating poleward along the coast from their equatorial source. These impose elevated sea levels,
a deeper thermocline, positive sea-surface temperature (SST) anomalies and either a decrease or
episodic reversal of coastal equatorward flow in the PCC and CCC (Figure 2) during the EN (warm)
phase. Decreased coastal sea levels, shallower thermoclines, colder SST conditions and strengthened
coastal equatorward flow occur during the La Niña (LN; cold) phase. Satellite data show that EN
anomalies can extend south of 40°S to include the entire coast of South America (Carr et al. 2002,
Strub & James 2002), but anomalies decrease with increasing latitude (Strub & James 2002,
Montecinos et al. 2003, Escribano et al. 2004a). EN conditions can also be imposed by local
anomalous winds, likely modulated via atmospheric teleconnections (Shaffer et al. 1999). Oceanic
ENSO signals originate as eastward-propagating equatorial Kelvin waves that propagate poleward
as coastally trapped waves (CTWs). Alongshore wind stress and offshore Ekman transport persist,
or even strengthen during EN periods (Carr et al. 2002, Escribano et al. 2004a), but upwell warm,
nutrient-poor water from above the deepened thermocline.
Mesoscale variability is evident as eddies and filaments that advect cold, high-pigment coastal
water for hundreds of kilometres offshore, often originating at headlands (Thomas 1999, Sobarzo &
Figueroa 2001, Mesías et al. 2003). Along the coast of central Chile, a transition zone of elevated
eddy kinetic energy is observed (Hormazábal et al. 2004). A wide zone (800–1000 km) of relatively
diffuse and low eddy kinetic energy is observed north of ~30°S where winds are weak but are
persistently favourable to upwelling. Eddy kinetic energy is stronger and more closely associated
with the coast (~600 km) in the region between ~30°S and 38°S where the winds, though stronger,
are more variable (Hormazábal et al. 2004). Away from the coast (>300 km), altimeter and drifter
data show two regions of maximum eddy kinetic energy, one along the Peru coast and another
centred at ~30°S off Chile. This is consistent with patterns of surface temperature and satellite-
measured ocean colour that show cold waters with high pigment concentrations associated with the
upwelling extending furthest offshore off Peru and off central Chile (Montecino et al. 2005), while
being restricted to an extremely narrow coastal band off northern Chile (Morales et al. 1999, Thomas
et al. 2001b). CTWs propagate poleward along the entire Peru and Chile coastlines, traceable to wind
fluctuations in equatorial regions (Hormazábal et al. 2001). CTWs raise and lower the pycnocline/
nutricline, influencing the effectiveness of upwelling, with dominant frequencies of days to weeks
off Peru (Enfield et al. 1987) and ~50-day periods off northern and central Chile (Shaffer et al.
1997, Hormazábal et al. 2001, Rutllant et al. 2004b). Ramos et al. (2006) show that variability of
equatorial origin at both annual and semi-annual periods impose strong modulation on isotherm
depth along the coast. CTWs appear especially energetic during EN periods and weaker during
LN periods and austral winter (Shaffer et al. 1999). At the shorter time-/space scales, diurnal cycles
in wind stress are important contributors to forcing along the arid northern Chilean coast, especially
in summer (Rutllant et al. 1998), but become less important with increasing latitude, where
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storm-mediated variability on 3- to 7-day cycles increases (Strub et al. 1998), with a maximum in
austral winter.
Coastal oceanography
Coastal waters have been defined in many different ways depending upon the reason (e.g., scientific,
geopolitical, international conventions, etc.) for their usage. Bowden (1983) defines them, from an
oceanographic perspective, as “those on the continental shelf and the adjoining seas”. If such a
definition would be used for the HCS, then the coastal strip would be rather narrow. Bottom depth
in some areas of the HCS (i.e., Antofagasta) goes from less than 100 m at distances of 10 km from
shore to more than 1000 m at 30 km. However, many characteristics of the plankton assemblages
at those distances show that they are still largely influenced by the proximity of the coast. Thus,
for the purposes of this analysis an alternative definition for coastal ocean is used: that part of the
ocean where the proximity of the continents affects the circulation and ecological processes. This
definition then includes coastal waters as defined by Bowden and the coastal transition zone defined
by Hormazábal et al. (2004).
The analysis of any oceanographic system is scale dependent (Haury et al. 1978, Rutllant &
Montecino 2002). Thus, if space–time is considered as a continuum, then isolating parts of it is an
observer decision, which may be influenced by its epistemological background (Ramírez 2005a)
and the characteristics of the ecosystem to be studied (Marín 2000). Haury et al. (1978), in a
classical article on scales of analysis in oceanography, propose two terms (mesoscale and gross
scale) to refer to those scales where the effects of flow structures such as filaments, squirts, meanders
and eddies (1–102 km) are dominant and where ecosystem patterns are advective (influenced by
the physics of the system) and biological. Coastal upwelling is one of the main mesoscale ocean-
ographic processes affecting the dynamics and the spatial and temporal structure of coastal eco-
systems along the EBCs (Strub et al. 1998, Montecino et al. 2005). Upwelling flow structures such
as filaments (Sobarzo & Figueroa 2001), squirts (Marín et al. 2003a, Marín & Delgado 2007) and
shadows (Castilla et al. 2002a, Marín et al. 2003b) have been described for the Chilean coast.
Herein, information about those structures is succinctly reviewed and the potential effects on the
ecology of the coastal ocean in the HCS are discussed.
The HCS, from the standpoint of coastal wind forcing, can be divided in two latitudinal areas
near 26°S (Figueroa 2002). From 26°S to the north, meridional, upwelling-favourable winds are
rather constant throughout the year; south of this latitude greater seasonality is observed. One
important upwelling focus in the northern zone is the Mejillones Peninsula (23°S). Observational
(Marín et al. 1993, Escribano et al. 2000, Marín et al. 2001, Olivares 2001, Sobarzo & Figueroa
2001, Escribano et al. 2002, Rojas et al. 2002, Marín et al. 2003b) and modelling studies (Escribano
et al. 2004b) have shown that the dynamics of the coastal ecosystems in that area largely depend
on the generation of upwelling filaments at Mejillones Peninsula. Indeed, the generation of filaments
in the northern tip of the peninsula (Punta Angamos) has been identified as the main mechanism
of nutrient enrichment in the surface layers (Marín & Olivares 1999). Furthermore, ‘upwelling
shadows’ within Mejillones Bay, an equator-facing bay located in the northern end of the peninsula,
have been dynamically linked to the generation of bifurcated filaments at Punta Angamos (Marín
et al. 2003b). This shadow is an important physical structure within the bay, affecting PP (Marín et al.
2003b) and the retention of planktonic organisms (Olivares 2001). An alternative mechanism,
described as an ‘upwelling trap’ by Castilla et al. (2002a) and also related to the coastal upwelling
dynamics, generates higher temperatures inside Antofagasta Bay, a pole-facing bay at the southern
end of the peninsula. In this case also the physically generated structure contributes to the retention
of planktonic organisms. Thus, mesoscale flow features (upwelling shadows and upwelling traps),
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associated with cold-water filaments, seem to play an important role not only in relation to the
biological productivity of coastal upwelling regions but also as mechanisms for the retention of
coastal planktonic species. If account is taken of the fact that Mejillones Peninsula is located in
the area where upwelling-favourable winds occur year-round, then those areas may constitute
recurrent retention zones.
Farther south, within the HCS area where upwelling is more seasonal, the Coquimbo Bay
System (30°S) is another important coastal upwelling centre. This system is located within two
coastal points (Punta Lengua de Vaca and Punta Pájaros) and is the site of intensive fisheries and
eco-tourism (Ramírez 2005b). Filaments, including bifurcated upwelling filaments (Moraga et al.
2001), are known to generate at Punta Lengua de Vaca, contributing with cold, nutrient-high waters
to the coastal system (Montecino & Quiroz 2000, Montecino et al. 2005). A recent Lagrangian
study conducted within the bay (Marín & Delgado 2007) shows that the dominant equatorward
flow is modulated at quasi-inertial frequencies, enhancing the coastal retention times. Furthermore,
Marín et al. (2003a) have shown that cold-water squirts generate at the northern end of the bay
system (near Punta Pájaros). Squirts seem to be geographically anchored to locations meeting the
requirements for their generation (Strub et al. 1991). Thus, it is highly likely that indeed the squirts
observed in the vicinity of Punta Pájaros are normally generated within the same area, making
them a recurrent mesoscale flow feature. As a test of this idea a simple model was built using the
ROMS model (Shchepetkin & McWilliams 2005) and initialised with the ROMSTOOLS package
(Penven 2003). The study area was defined by longitudinal boundaries set at 74°W and 70°W and
latitudinal boundaries at 32°S and 24°S (see also Figure 3). This model has been run for 2 yr in
other eastern boundary systems such as the California Current System (Marchesiello et al. 2003)
to obtain a statistical equilibrium condition. However, in the present test case a squirt was generated
just after 17 days of integration, which closely resembles the squirt observed in an SST image
obtained in January 2005 in shape, size and location (Figure 3). The interesting, and indeed
serendipitous, observation resulting from this numerical exercise is that transient modes developed
as perturbations (e.g., baroclinic instabilities) from climatological mean conditions generate coastal
flow structures (squirts) which are recurrently found within the HCS. Satellite observations and
Lagrangian drifter data (Marín & Delgado 2007) have in fact shown that this squirt is a recurrent
feature in the area, reaching distances on the order of 140 km offshore. Squirt speeds, estimated
both through feature-tracking analysis (Marín et al. 2003a) and Lagrangian drifters (Marín &
Delgado 2007), range between 0.2 and 0.3 m s−1. Thus, considering that the lifetime of a single
squirt is related to the active period of equatorward wind events, which for the area range between
3 and 7 days (Rutllant et al. 2004a), coastal organisms trapped within the squirt are likely to reach
100–200 km offshore in a period of less than a week.
The conclusions that may be offered as a result of this brief, and highly condensed, analysis
of the prevailing mesoscale coastal features of the HCS are nevertheless far reaching. In the first
place, if these features are intrinsic to the HCS (generated as a result of coastline geometry, bottom
topography and prevailing flow conditions including perturbations) and not dependent on large-
scale, low-frequency forcing (e.g., ENSO), then a whole new array of multiscale (nested) models
are necessary to generate predictable bio-oceanographic coastal patterns for the HCS. Second,
coastal upwelling areas can be described as a dynamic mosaic of nearshore retention/offshore
expatriation patches or sectors. Thus, for a planktonic organism, remaining in a specific location
(i.e., local populations) within the HCS coastal zone becomes a probabilistic process. If, for
example, larvae are entrained within an upwelling shadow then there is a high chance that they
will remain in the same sector for a period close to a week. If, on the other hand, the larvae are
entrained within a squirt, then in a period close to 24 h they will be expatriated offshore. However,
since the seascape is dynamic and depends upon the upwelling condition, it is not possible to ensure
that a given geographic location will always act as a retention or expatriation locality.
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26°S
73° 72° 71°
28°S 28° 28°
30°S
29° 29°
73°W 72°W 71°W 30°

30°
73° 72° 71°

60 0 60 120 180 Kilometers
Figure 3 (See also Colour Figure 3 in the insert following page 344.) Result of the squirt simulation by means
of the Regional Oceanic Modeling System (coloured image) for 17 January in comparison to the thermal
squirt (NOAA satellite, black-and-white image) observed on 15 January 2005. Temperature coding of the
coloured image goes from blue (lower temperature) to red (higher temperature) while for the NOAA image
it goes from lighter to darker tones of grey. White areas in the NOAA image correspond to clouds. The study
area in the model was defined by longitudinal boundaries set at 74°W and 70°W and latitudinal boundaries
at 32°S and 24°S. The resolution was 1/10°, approximately 10 km, with 20 vertical levels, a minimum depth
of 50 m and a smoothed bathymetry. The baroclinic time step was set to 900 s. Surface forcing (wind stress,
heat fluxes, freshwater flux, SST, sea-surface salinity, and short-wave radiation) was derived from the monthly
climatology found in the COADS dataset (Da Silva et al. 1994) and interpolated into the model grid. The
temperature and salinity initial conditions for the model were obtained for the month of January from the
‘World Ocean Atlas 2001’ (Conkright et al. 2002). The Ekman and geostrophic velocities at the boundary
were obtained combining these data with COADS winds, following Marchesiello et al. (2001), with level of
no motion defined at 500 m (Marchesiello et al. 2003).
The water column chemistry

The northern Chilean ocean margin (~18–30°S) presents distinctive characteristics in topography,
climatic and oceanographic conditions, which modulate PP and water column chemistry. It features
a comparatively low PP (for the HCS), and despite semi-permanent wind-driven upwelling some
areas are considered as high-nutrient low-chlorophyll (HNLC) environments (Torres 1995, Daneri
et al. 2000). These observations are in contrast to the localised prominent upwelling cells with
relatively high PP, such as off Iquique (21°S), Antofagasta (23°S) and Coquimbo (30°S) (0.5–9.3 g
C m−2 d−1; González et al. 1998, Daneri et al. 2000, Thomas et al. 2001a).
Studies carried out in the main upwelling centres indicate that highest PP foci occur close to
the coast over the narrow continental shelf and fuel major fisheries, with catches that represent
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40% of the annual landings of the HCS (Escribano et al. 2003 and references therein). This
production also constitutes an important way of sequestering CO2 and supports a high rate of
particulate organic matter (POM) exported to depth (González et al. 2000a, Pantoja et al. 2004).
This material, which is partly remineralised in the water column, strengthens the oxygen-minimum
zone (OMZ) and promotes biogeochemical anaerobic processes. In this sense, a sequence of
mechanisms that are determined by the oceanographic conditions is regulating the chemistry of the
water column and the sea bottom. These processes gain relevance in several aspects of material
exchange (i.e., gas fluxes as CO2 and N2O), implying that this area could play a key role in the
main global cycles (i.e., oceanic productivity, global warming, authigenic carbonatic and phospho-
rite mineral formation, etc.).
Oxygen distribution and relevance in organic carbon remineralisation

Most research efforts have focused on specific areas where high productivity cells are recognised,
but the interactions with biogeochemical processes are still poorly understood. Along the northern
margin of Chile, a well-developed OMZ is observed between 100 and 500 m water depth (Blanco
et al. 2001). This zone is basically a mid-water feature since the topography of the margin precludes
the OMZ to impinge on a large area of the bottom, except off Antofagasta (Mejillones) where it
touches the bottom from 50 m to 300 m depth. Normally the shelf is extremely narrow from
southern Peru to northern Chile (10–15 km) in comparison with central Peru and southern-central
Chile (40–60 km; Strub et al. 1998), where the OMZ extends over a wide area of the shelf, promoting
distinct biogeochemical processes (Gutiérrez 2000, Neira et al. 2001). This characteristic of northern
areas could thus affect pathways (i.e., aerobic or anaerobic) associated with organic matter (OM)
degradation in the sediments, which is an important source of regenerated nutrients to the water
column. Off Mejillones, a high percentage (86%) of photosynthetically produced particulated
protein is being degraded within the upper 30 m of the water column (Pantoja et al. 2004), coinciding
with oxygenated waters. In consequence, OM reaching the sediments at greater depths is depleted
of proteins. Over the shallower shelf sediments, where also preserved fish debris and bones are
found (Milessi et al. 2005), high pigment concentrations have been reported (42–100 µg g−1 of
chloroplastic pigment equivalents in surface sediments; P. Muñoz et al. 2004a, 2005). Thus, there
is a narrow band of inshore sediments that are enriched in fresh OM coming from the water column.
The remineralisation of this material could generate an important flux of nutrients contributing to
fertilisation of the water column. Similar predictions can be made for other areas of high PP along
the coast of northern Chile, where upwelled waters containing preformed nutrients are enriched
with recycled nutrients derived from the OM degradation in shelf sediments. The relevance of the
sea floor in water column fertilisation and biological productivity as well as its relevance in the
global carbon cycle has not been well examined. Some information is available for the role of
sediments near the main upwelling centres, but nothing is known about the biogeochemical pro-
cesses along the large extent of the margin between the main upwelling centres mentioned here.
The high OM degradation rates result in CO2 supersaturated waters that, in combination with
the CO2 input from upwelled waters, favour the CO2 flux from the ocean to the atmosphere. Some
studies off Antofagasta (23–24°S) and Coquimbo (30°S) have measured a saturation of >200% in
upwelled waters, an f CO2 up to 1000 µatm and CO2 flux ~3.9–4.0 mol C m−2 yr−1 (Torres et al.
2003 and references therein). These authors concluded that CO2 outflux is a highly variable short-
term process, depending on the intensity of the wind-driven upwelling, the depth of the upwelled
waters, the OM degradation (positively correlated with the apparent oxygen utilisation, AOU) and
the biological uptake. Furthermore, the high degradation rates of organic carbon by the microplank-
ton community (dissolved organic carbon-, DOC; 1.1–21.6 µM h−1; G. Daneri unpublished data)
and high respiration rates (81–481 mmol O2 m−2 d−1; R.A. Quiñones unpublished data) indicate
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that nutrient recycling in the water column is an important process in the CO2 production, resulting
in high concentrations of CO2 and outgassing when upwelled waters are warming up at the sea
surface. A substantial proportion of the carbon assimilated by primary producers is reaching the
bottoms via biogenic CaCO3 flux, as has been observed off Coquimbo (30°S) where sediment traps
located at 2300 m water depth (~180 km off the coast) revealed that almost 40–90% of carbonate
flux is associated with some species of foraminiferans characteristic of upwelling areas (H.E.
González et al. 2004a, Marchant et al. 2004). These authors suggest that biogenic CaCO3 is the
main pathway by which carbon is removed from the upper ocean, controlled by autochthonous and
allochthonous foraminiferans, that is, large-size organisms with high sinking rates (1.5 days) and
smaller organisms that are laterally advected. Therefore, part of this carbon is exported offshore,
sequestered from the water column and preserved in the sediments (Hebbeln et al. 2000a,b), but it
has not been clearly established what percentage of the total CO2 assimilated by primary producers
this CaCO3 flux represents.
Macro- and micronutrient distribution

The distribution of nutrients shows a high variability in the water column associated with upwelling
pulses and mixing. High concentrations occur inshore and usually decrease in the offshore direction,
followed by decreasing pigment concentrations (Escribano et al. 2003, Marín et al. 2003a, 2004a,
Peñalver 2004). Off 30°S, high surface concentrations of nitrate, phosphate and silicate have been
reported (~5–15, 0.5–1 and 5–8 µM, respectively; Peñalver 2004). The nutrient distribution at this
latitude is also affected by the topography of the area, where several islands reduce the circulation
and mixing of the water column (Peñalver 2004). In northern Chile, between 20°S and 22°S, high
concentrations of nitrate were also observed near shore at <100 m water depth (0–20 µM; Escribano
et al. 2004b, Pantoja et al. 2004, Herrera & Escribano 2006). In general, the distribution of nitrate
showed low subsurface concentrations (Pantoja et al. 2004, Peñalver 2004) associated with high
nitrite concentrations (>2 µM; Morales et al. 1996) and coincident with low oxygen concentrations
(<0.5 ml L−1). This implies that denitrification is an important process in the nitrogen recycling, as
has been recognised for other OMZ regions (Codispoti & Christensen 1985, Codispoti et al. 1986,
Cornejo et al. 2006).
Another important aspect that controls the biogeochemistry of the water column is the hydro-
logical regime. Northern Chile, with the Atacama Desert, is an extremely arid zone without relevant
fluvial inputs into the ocean. The maximum amount of precipitation is ~0.5–78 mm yr−1, which is
significantly lower than in central and southern Chile (300 to >1000 mm yr−1; www.meteochile.cl).
In general, rivers play an important role in the fluxes of trace metals, nutrient and particulate matter
to coastal waters, and some of these components are considered to be important factors determining
PP in the water column. For example, Fe has been proposed as a factor limiting PP in waters
enriched with other macronutrients but low in pigment concentrations (Martin & Gordon 1988,
Martin et al. 1993, Coale et al. 1998). Some recent studies suggest that this element should be
relevant in PP off northern Chile, where low dissolved Fe concentrations have been measured
(0.6–1.4 nM; R. Torres unpublished data). Additionally, other elements such as Cd and Co may
also play an important role in biological processes. The concentration of dissolved Co in the water
column shows a similar vertical distribution as macronutrients in an upwelling region off Peru
(8–10°S), apparently controlled by biological uptake and remineralisation (Saito 2005). In the same
sense, dissolved Cd in the water column for Mejillones Bay shows a typical micronutrient-like
distribution (23°S; J. Valdés unpublished data). Low values of dissolved Cd (~0.4–1.6 nM) were
found in the water column, while very high concentrations were measured in surface sediments
(~60 µg g–1; Valdés et al. 2003). In other coastal waters of northern-central Chile, high Cd values
in sediments are probably associated with biological uptake and subsequent deposition in the
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sediments (~30°S; <6.6 µg g−1; P. Muñoz unpublished data). Therefore the uptake of trace elements
by primary producers and subsequent export to sediments appears to be another important factor
controlling the concentrations of metals and nutrients in the water column.
Dominant primary producers and their role

in the pelagic food web
The HCS off Chile demonstrates high PP associated with wind-driven upwelling events of different
intensities and frequencies along the South American coast (Strub et al. 1998). In northern Chile
(21–23°S, off Iquique and Antofagasta), a permanent upwelling sustains high PP throughout the
entire year (3–9 g C m−2 d−1; Daneri et al. 2000, Montecino & Quiroz 2000, Iriarte & González
2004) while in the central-southern area (i.e., Concepción at 36°S) the upwelling events are mainly
concentrated in the spring–summer period (Strub et al. 1998). In the above coastal areas intense
upwelling of nutrient-replete waters fertilise the euphotic zone (e.g., NO3−: 10–25 µM) (Escribano
et al. 2004b) creating suitable conditions for large-size primary producers, and in turn causing
profound effects on the water column processes such as carbon flow through the pelagic food web
and carbon export toward deeper layers of the ocean.
Off northern Chile a weak seasonality in chlorophyll-a (chl-a) concentration showed slightly
higher values during the winter and early spring, whereas in central southern Chile (30–40°S), the
annual maximum in chl-a concentration occurred during the summer (Thomas et al. 2001a). In the
northern coastal area off Chile (18–24°S) the highest chl-a concentrations (15–20 mg m−3) were
mostly restricted to a narrow inshore zone (<37 km) and were associated mostly with microphyto-
plankton (>20 µm), whereas pico- and nanophytoplankton (pico: 0.7–2.0 µm; nano: 2.0–20 µm)
predominate in the offshore, mostly oligotrophic zone (Morales et al. 1996, Iriarte & González
2004). Off central Chile (33–37°S) upwelling-favourable south-southwest winds predominate dur-
ing the austral spring and summer months when chl-a ranged between 3.8 and 26 mg m−3 compared
with 1.0–2.5 mg m−3 recorded during winter (González et al. 1989, Montecino et al. 2004).
The permanent and seasonal upwelling off Antofagasta (Mejillones Bay) and Concepción
(Concepción Bay), respectively, produce a highly productive phytoplankton assemblage, dominated
by no more than 10 species of chain-forming diatoms, such as Chaetoceros spp., Thalassiosira
spp., Guinardia delicatula, Rhizosolenia spp., Detonula pumila, and Eucampia cornuta (Rodríguez
et al. 1996). The cell sizes of the phytoplankton species along the northern coast of Chile generally
are >20 µm (Morales et al. 1996, Iriarte & González 2004), and high microphytoplankton abun-
dances in the sediment record correlate positively with intense/more frequent upwelling events
(higher PP) in waters of Mejillones Bay (Ortlieb et al. 2000). In contrast, small-size phytoplankton
(<20 µm) predominate in oligotrophic regimes, associated with the intrusion of subtropical, nutrient-
depleted, warmer subtropical waters (STW) (Iriarte et al. 2000, Iriarte & González 2004). In such
cases, small dinoflagellates (Gymnodinium spp., 5–25 µm) and autotrophic flagellates dominated
the phytoplankton assemblages (Iriarte et al. 2000). In Concepción Bay (37°S), few numerically
dominant diatom species represented >80% of the total diatom abundance during upwelling events
(González et al. 1987), and in correspondence with this, the highest biomass was concentrated in
the microphytoplankton fraction (>20 µm) from winter through spring (González et al. 1989).
The first studies that considered several trophic levels of the pelagic system (Peterson et al.
1988) and trophic models of carbon flux (Bernal et al. 1989) were conducted in the coastal area
off Concepción during the late 1980s. These original studies supported the classical view that the
upwelling areas are characterised by short food chains dominated by large chain-forming diatoms
and few small clupeiform fish species or the ‘traditional food chain’ (Ryther 1969). This view has
recently been challenged, highlighting the relevance of the microbial loop (Troncoso et al. 2003,
210
Vargas & González 2004) and gelatinous food web (H.E. González et al. 2004b). These trophic
flows are important throughout the whole year in oceanic areas and are highly relevant during the
non-productive periods (including EN events) in coastal upwelling areas.
The carbon budget of the photosynthetically generated OM in the coastal areas of the HCS has
been under debate for many years (Bernal et al. 1989, González et al. 1998). It is accepted nowadays
that the fraction of the PP removed from the photic zone, which is highly variable on an annual
basis (Hebbeln et al. 2000a), strongly depends on the various biological (internal metabolism),
physical (stratification/mixing), and chemical (nutrient rich/poor waters) processes involved as well
as the time of year. However, the sources of this variability, both in space and time, have been
poorly analysed until recently (Morales & Lange 2004), mainly because of the lack of long-term
time-series studies. In Figure 4 the main pathways of the photosynthetically generated organic
carbon are depicted. Very high bacterial secondary production (BSP) has been reported in the
coastal area of northern-central Chile (Troncoso et al. 2003, Cuevas et al. 2004), suggesting a tight
coupling between PP and BSP. The pivotal role of bacteria is supported by the exceptionally high
Primary production
5700 Zooplankton
grazing
Microplankton
respiration 455
Iquique −20°
1513
Bacterial secondary
production
1106 Antofagasta
−25°
Export
114
production
A Coastal area off Antofagasta Southern latitude
Coquimbo −30°
Primary production
4570 Zooplankton
grazing
Microplankton
respiration 321 −35°
1400
Bacterial secondary Concepción
production
2300
−40°
Export 336
production
B Coastal area off Concepción
Figure 4 Photosynthetically generated carbon and its flows (rate estimates in mgC m–2 d–1) through the more
relevant biological processes in the upwelling system off Antofagasta (23°S) and Concepción (37°S).
211
MARTIN THIEL ET AL.
degradation rates of dissolved organic carbon in Concepción Bay (1–21 µM h−1; G. Daneri unpub-
lished data) and the high BSP rates (range of 1100–2300 mg C m−2 d−1 or 19–50% of PP). Microbial
community respiration rates (Eissler & Quiñones 1999) are also very high along the HCS, reaching
average values of 1450 mg C m−2 d−1 (~28% of PP). Finally, both zooplankton grazing and export
production (González et al. 2000b, Grünewald et al. 2002) gave values between 100 and 500 mg
C m−2 d−1 (or mean values between 2% and 10% of PP). These carbon flows are more representative
of the coastal upwelling systems of Antofagasta and Concepción because they are the most studied
areas (from an oceanographic point of view) along the Chilean coast. Estimations of PP for the
HCS along the Chilean coast are similar to those of the Peru (4000 mg C m−2 d−1; Walsh 1981)
and about 2-fold higher than those of the California (1000–2500 mg C m−2 d−1; Olivieri & Chavez
2000) upwelling systems. In addition, typical upwelling values for the flow of OM through bacteria
in the HCS (19–50% of PP) are well within the range (3–55% of PP) of those described for other
upwelling systems in the world oceans (Ducklow 2000).
Processes affecting primary production and export processes

In coastal areas of central and southern Chile there is a distinctive seasonal pattern involving the
development of maxima (spring) and minima (winter) in phytoplankton biomass and PP during an
annual cycle (Ahumada 1989, González et al. 1989). In contrast, high and more constant phyto-
planktonic biomass and PP has been observed during an annual cycle along the northern coast off
Chile (Marín et al. 1993, Rodríguez et al. 1996, Marín & Olivares 1999). Among the factors that
might control the PP, light limitation (P. Montero & G. Daneri unpublished data) and Fe availability
(Hutchins et al. 2002, R. Torres unpublished data) have been suggested for the Concepción and
Coquimbo upwelling systems, respectively. In addition, high microzooplankton grazing might
control the PP during non-upwelling conditions (i.e., winter) in Concepción Bay upwelling (Böttjer
& Morales 2005).
The understanding of the factors that regulate the magnitude and the variability of phytoplankton
PP is quite complex in the HCS due to the geographically distinctive upwelling areas (wind stress,
topography), seasonal changes (winter vs. spring) and coastal–oceanic gradients. Integrated values
in Table 1 indicate that the variation in chlorophyll concentration is coherent with changes in PP;
that is, estimates are lowest in the Coquimbo upwelling system and highest in Antofagasta and
Concepción coastal upwelling areas. Higher chlorophyll-specific productivity at Coquimbo and
Antofagasta during EN 1997–1998 (2.5–2.8 vs. 1.0–2.0) indicates that the increase in specific
productivity did not result solely from a biomass decrease, but from a change in the phytoplankton
size distribution (therefore in species composition), from the larger size class (microphytoplankton)
to smaller size classes (pico- and nanoplankton). The intrusion of oligotrophic oceanic waters into
the coastal area off Coquimbo (Shaffer et al. 1995) and Antofagasta (Iriarte & González 2004)
during an EN could be a possible explanation for the low productivity and the dominance of smaller
phytoplankton size fractions and their large contribution to total PP. This feature suggests that
biological and physiological shifts occur at the phytoplankton species level in order to counteract
the change in prevailing physical and chemical conditions in those areas (Montecino & Quiroz
2000, Pizarro et al. 2002). On the other hand, in the permanent/seasonal coastal upwelling of cold
nutrient-rich waters in enclosed areas like Concepción Bay and Mejillones Bay, PP estimates
increase up to 12 g C m−2 d−1. In the above-mentioned areas, the microphytoplankton fraction
accounted for >50% for the total autotrophic biomass and PP. In general, the range of specific rate
of productivity in the three upwelling areas could indicate that physiological factors such as nutrient
supply and/or light availability may regulate the seasonal signal of productivity in those areas,
whereas top-down processes such as grazing and production export might be important in removing
a fraction of the generated photosynthetic carbon (Figure 4, Table 1). Furthermore, high biological
212
Table 1 Range of primary productivity, chlorophyll, chlorophyll specific primary productivity rate and main phytoplankton taxa between the
22° to 37°S Humboldt Current System. Primary productivity and chlorophyll estimates are integrated to the 1% light penetration depth
PB Dominant size
PP Chl. a (mmol C mg class (as % of
Study area (mg C m–2 d–1) (mg m–2) Chl. a–1 d–1) the total Chl. a) Main phytoplankton taxa Reference
Antofagasta 338–6063 11.7–175.4 2.8 48–68% Gymnodinium sp., Pseudo-nitzschia cf. delicatissima Pizarro et al. 2002
(22–2°S) nanoplankton Autotrophic flagellates Iriarte et al. 2000
El Niño 1997-1998 Ulloa et al. 2001
Troncoso et al. 2003
Antofagasta 1100–8100 47–695 1.0 60–86% micro- Chaetoceros spp., Thalassiosira spp., Rhisozolenia Iriarte & González 2004
(22–23°S) phytoplankton spp., Detonula pumila, Guinardia delicatula,
Non El Niño Eucampia cornuta

2000–2002
Coquimbo-Valparaíso Total: 140–2955 8.0–92.5 2.5 Winter: 59% Detonula pumila, Leptocylindrus danicus, Montecino et al. 1996
(30-33°S) Summer: 605–2224 nanoplankton Pseudo-nitzschia pseudoseriata, Montecino & Pizarro 1995
1992–1994, 1995 Winter: 602–1012 Prorocentrum micans Avaria & Muñoz 1982
Montecino & Quiroz 2000
213
Troncoso et al. 2003
Humboldt Current 53–141 Winter: >50% Coast: Pseudo-nitzschia pseudoseriata, Chaetoceros Morales et al. 1996
System compressus, Leptocylindrus danicus, Rhizosolenia Avaria et al. 1982
(19–22°S) imbricata, Ceratium tripos, Diplopsalis lenticula
Spring: 65% Oceanic: Chaetoceros coarctatus, Ch. dadayi,
micro- Ceratium contortum, C. gibberum, C. macroceros,
phytoplankton Dinophysis rapa, Ornithocercus magnificus
Las Cruces (33°S) 1–13.5 Narvaez et al. 2004
1999–2000 (mg m–3)
Concepción Bay and Total: 200–21,000 Winter: 48% Thalassiosira spp., Detonula pumila, González et al. 1989
Gulf of Arauco Winter: 481–1600 nanoplankton Chaetoceros socialis, Chaetoceros curvisetus, Iriarte & Bernal 1990
(36–37°S) Spring: Spring: Spring: 2.0 Spring: 84% Skeletonema costatum, Leptocylindrus danicus, Ahumada 1989
1770–16,670 208–544 micro- Guinardia delicatula Daneri et al. 2000
phytoplankton Troncoso et al. 2003

Montecino et al. 2004
Cuevas et al. 2004
MARTIN THIEL ET AL.
productivity observed along the northern and central coast of the HCS (23–37°S) is probably
fertilising oceanic and oligotrophic areas through the formation of eddies and filaments that transport
coastal water offshore (Marín et al. 2003a, Hormazábal et al. 2004, Letelier et al. 2004). Finally,
much of what is known about upwelling dynamics, phytoplankton ecology and their role in
biogeochemical cycles along the Chilean coast has been mainly gathered from selected upwelling
areas (i.e., Antofagasta, Coquimbo, Concepción, Valparaíso) with scarce or no information from
areas in-between.
Zooplankton consumers
Zooplankton consumers are considered the link between primary producers and higher trophic
levels in the pelagic realm of the world ocean. This function also implies a role to regulate the rate
at which phytoplankton-C can be transferred through the food web or retained as zooplankton
biomass. This rate greatly depends on population turnover rates, which vary widely among zoo-
plankton taxa. Thus, the knowledge of the population biology of dominant species, the community
structure and its variation, become key issues in understanding the ecological and biogeochemical
role of zooplankton. In the HCS substantial progress has been made in the last decades about some
of these issues. In the following sections we summarise the major advances in understanding
population, community and ecosystem processes involving zooplankton in the upwelling region
off northern Chile. Much less progress is being made in molecular, genomic and genetic biology
of zooplankton in this region. However, the need for a better understanding of zooplankton ecology
in the region will most likely motivate the use of molecular tools in the near future.
Biogeographic and biodiversity issues

In the upwelling region off northern Chile most species or species assemblages are considered to
be part of the subantarctic fauna. This fauna originates in the Austral region, but it is then advected
northward. In northern Chile and off Peru, it mixes with some species of tropical and equatorial
origin. The most studied taxa are Copepoda (Heinrich 1973, Vidal 1976, Hidalgo & Escribano
2001) and Euphausiacea (Antezana 1978, D. Fernández et al. 2002). Nearly 60 species of copepods
have been identified, of which the dominant in the coastal zone are Calanus chilensis, Centropages
brachiatus, Paracalanus parvus, Acartia tonsa, Eucalanus inermis, Oithona similis, Oncaea
conifera and Corycaeus typicus. Calanus chilensis is an endemic species of the HCS (Marín et al.
1994), whereas Centropages brachiatus, Paracalanus parvus, Acartia tonsa and Oithona similis
are cosmopolites and widely distributed along the Chilean coast. Eucalanus inermis is a typical
tropical species, although the genus Eucalanus may contain about six species, which have not yet
been clearly defined. Among euphausiids, the most abundant and endemic species of the HCS is
Euphausia mucronata (Antezana 1978). This species is closely associated with upwelling centres
off northern Chile (Escribano et al. 2000) and performs an extensive vertical migration into the
OMZ. Euphausia eximia is another abundant species, which increases in number during EN
(Antezana 1978, González et al. 2000b). Gelatinous zooplankton, on the other hand, has received
much less attention, although it may form dense aggregations at times of the year in the coastal
upwelling zone (Pagès et al. 2001), and it can exert a strong predation pressure on copepods
(Giesecke & González 2004, H.E. González et al. 2004b, Pavez et al. 2006) and possibly on fish
larvae.
Species replacements seem to occur in the zooplankton associated with EN versus LN regimes
(Hidalgo & Escribano 2001). Dominant species alternate between copepods (mainly Calanus and
Eucalanus genera) and euphausiids during upwelling conditions and cyclopoid copepods, Paracalanus
214
and Acartia during EN events (González et al. 2002). Warm conditions during an EN may also
cause reduced size of copepods at maturity (Ulloa et al. 2001). All these changes in structure of
the pelagic system may have profound implications on the functioning and productivity of this
region (see, e.g., Alheit & Niquen 2004) and should also be considered in future ecosystem studies.
Zooplankton and coastal upwelling in northern Chile

In the coastal zone off northern Chile, wind-driven upwelling is the principal driver of biological
productivity. Pelagic organisms benefit by high productivity in upwelling sites. However, upwelling
zones comprise strongly variable environments and zooplankton must cope with changing condi-
tions in this zone. The understanding of these physical and biological interactions in the water
column of upwelling systems may give much insight into the key processes that control production
and biological diversity of pelagic assemblages. In northern Chile, examples of physical–biological
interactions taking place during coastal upwelling are restricted to particular conditions and sites
(e.g., Escribano & Hidalgo 2000a, Escribano et al. 2001, 2002, Giraldo et al. 2002). Other studies
(Marín et al. 2001) have shown some relevant findings that may help understanding mechanisms
through which pelagic populations are able to maintain their populations in the food-rich coastal
zone. Off Mejillones Peninsula (23°S), the interaction between a poleward flow and cold upwelling
plumes may generate large eddies by which non-migrant plankton can be maintained nearshore.
This type of circulation may act as a retention mechanism to avoid offshore advection of zoo-
plankton (Giraldo et al. 2002). Thus, most species maintain their population in the food-rich
upwelling centres being recirculated by near-surface currents. In other upwelling systems, such as
the Benguela Current, zooplankton is maintained nearshore by vertical migration behaviour
(Verheye et al. 1992, Roy 1998). Organisms advected offshore migrate to deep water and then are
returned to shore by a compensatory flow. In northern Chile, however, vertical migration does not
seem to be an important or widely used behaviour. In fact, most dominant species appear strongly
constrained to the upper layer (<100 m) (Escribano & Hidalgo 2000a) because of oxygen-depleted
waters in the OMZ, which come close to the surface (<50 m) during upwelling (Morales et al. 1999).
Upwelling in the coastal zone off northern Chile may exhibit much spatial variation character-
ised by discrete upwelling centres, such as those usually observed off Iquique (20°S), the river Loa
(21°S), Mejillones (23°S) and Coquimbo (30°S) (Fonseca & Farías 1987). Zooplankton populations
tend to aggregate in these upwelling centres and hence they are also very patchily distributed
(González & Marín 1998, Escribano & Hidalgo 2000a). Spatial heterogeneity in plankton distri-
bution linked to variability in oceanographic conditions, such as temperature and food quantity and
quality, may be one of the major causes of variability in population growth and secondary production
along the coast off northern Chile (Giraldo et al. 2002).
Zooplankton and the oxygen minimum zone

The OMZ system is a prominent feature in northern Chile closely related to the ESSW. This oxygen-
depleted water mass is also strongly linked to wind-driven upwelling in the coastal area. The ESSW,
which normally occupies the intermediate (200- to 500-m) layer (Blanco et al. 2001), may ascend
to shallow waters (<50 m) near the coast due to upwelling (Morales et al. 1999). The influence of
this low-oxygen water on pelagic communities of the coastal zone is not well understood. Dominant
zooplankton, which usually aggregates near the upwelling centres in this region (Escribano &
Hidalgo 2000b), must cope with such low-oxygen conditions. The options are either avoiding the
OMZ and restricting the population to the upper oxygenated layer, as earlier reported for some
dominant copepods (e.g., Escribano 1998), or evolving some metabolic adaptations to withstand
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MARTIN THIEL ET AL.
poor oxygen conditions, such as those described in González & Quiñones (2002). It has been
observed that several abundant epipelagic species concentrate in the upper 50 m without exhibiting
diel vertical migration (DVM) (Escribano 1998, Escribano & Hidalgo 2000a), although some
euphausiids, such as Euphausia mucronata, may temporarily enter the OMZ (Antezana 2002), or
some others like the copepod Eucalanus inermis may even reside in it (Hidalgo et al. 2005a). Thus,
the OMZ cannot be considered as just a constraint for vertical distribution because several species
may use it as their habitat, either temporarily or permanently. The ecological and biogeochemical
consequences of entering and living within the OMZ should be considered as relevant issues. Future
studies of species life cycles and behavioural or metabolic adaptations may provide novel findings
for life under low oxygen, as described for benthic organisms inhabiting these systems (Helly &
Levin 2004). Cycling and vertical fluxes of C and N mediated by zooplankton migration and vertical
distribution may be substantially modified by the low-oxygen and highly reduced environment.
These issues have received little attention for marine zooplankton associated with OMZ systems
(e.g., Wishner et al. 1998).
In the upwelling region off northern Chile, the most abundant species are usually closely related
to coastal upwelling plumes (Escribano et al. 2000, Giraldo et al. 2002). These species have been
well identified (Heinrich 1973, Hidalgo & Escribano 2001). Among dominant ones, the studies of
horizontal and vertical distribution have been focused on the calanoids Calanus chilensis (Escribano
1998), Centropages brachiatus (González & Marín 1998) and Eucalanus inermis (Hidalgo et al.
2005a) and on the euphausiid Euphausia mucronata (Escribano et al. 2000, Antezana 2002). The
available information indicates that Calanus chilensis and Centropages brachiatus are mostly
restricted to the upper layer without performing substantial DVM (Escribano 1998, Escribano &
Hidalgo 2000a). By contrast, Eucalanus inermis, the dominant species among a complex of four
to five species of the genus Eucalanus that coexist in this region, may remain in the upper boundary
of the OMZ with limited excursion into surface waters (Hidalgo et al. 2005a). Meantime, Euphausia
mucronata has been suggested as actively and daily migrating into the OMZ (Antezana 2002). A
summary of the vertical extent of species habitats for zooplankton in this region has been recently
constructed from several cruises (Escribano et al. accepted) and is illustrated in Figure 5. Zoo-
plankton can indeed occupy the entire water column despite the presence of an intense OMZ.
Occurrence and vertical movements of various species may ensure a substantial contribution of
zooplankton to the vertical export of OM.
Zooplankton life cycles and population dynamics

Life cycles and population dynamics are certainly important issues for understanding the bio-
geochemical and ecological role of zooplankton in this region. When examining annual life cycles
there are some important factors to consider. Since the entire upwelling region is subjected to
interannual variability due to the ENSO cycle, different years might induce changes in populations.
Water column warming, an abrupt thermocline and oxycline deepening characterises the EN
conditions in northern Chile (Ulloa et al. 2001, Escribano et al. 2004a), in contrast to a typically
shallow thermocline and OMZ indicating a ‘normal’ (LN condition) upwelling situation. However,
despite oceanographic variability some species, such as Calanus chilensis, seem to have a rather
stable annual cycle from year to year independent of ENSO variation (Escribano & Hidalgo 2000b).
Dominant zooplankton species have been suggested to be strongly associated with upwelling centres
(González & Marín 1998, Escribano & Hidalgo 2000b), and they can thus complete their life cycles
within the upwelling zone, growing at temperature-dependent rates under non-limiting conditions
of food (Escribano & McLaren 1999, Giraldo et al. 2002). Lack of food shortage has even been
suggested during EN conditions (Ulloa et al. 2001). Continuous reproduction and multiple gener-
ations throughout seasons characterise life cycles of copepods in this region (Escribano & Rodríguez
216
0
20
40
60
80
100
120
140
160
180
Depth (m)
200
220
240
260
Centropages brachiatus (1)

280
Pleuromamma gracilis (3)
Euphausia distinguenda
Scolecithrichella bradyi ?
Paracalanus parvus (1)
Nematoscelis megalops
300
Euphausia mucronata
Euphausia tenera
Eucalanus attenuatus
Eucalanus inermis (1)

Subeucalanus crassus
Euphausia eximia (3)

Corycaeus typicus (1)
Nematoscelis flexipes
Calanus chilensis (1)
Eucalanus subtenuis
Oncaea conifera (2)
320
Euphausia recurva
Stylocheiron affinis
Candacia rostrata
?
Euaetideus bradyi
Acartia tonsa (2)
Oithona similis
340
Euphausia sp.
Candacia sp.
Euchaeta sp.
360
380
400
420
Figure 5 Dominant habitat and daily movements of zooplankton species in and out of the main core of the
oxygen minimum zone (OMZ) (shaded area) during coastal upwelling at northern Chile. Day (white dots)
and night (black dots) positions represent the depth distribution of each one of the species listed and the arrow
the extent of their vertical movement. These positions were estimated from depth-weighed averages of
abundances. Data are from MinOx cruise carried out in March 2000 (modified from Escribano et al. accepted).
(1) Dominant species found in upwelling centers, (2) inshore species, (3) species associated with El Niño
events.
1994, Hidalgo et al. 2005b). However, strong variation in population size can be observed at some
times of the year (Escribano 1998, Ulloa et al. 2001, Hidalgo et al. 2005b). The populations of
annual species typically show a sudden collapse that tends to occur by the end of the summer. This
pattern has been described for Eucalanus inermis (Hidalgo et al. 2005b), Calanus chilensis and
Centropages brachiatus (Hidalgo & Escribano submitted) and Euphausia mucronata (Escribano
et al. accepted). The conceptual model of population dynamics for these species (Figure 6) considers
a two-stage population, which may grow exponentially during the spring and early summer at low
mortality rate and then exhibit an abrupt decay at high mortality rate due to increased predation.
In this model the increase in predation pressure coincides with the rise of the OMZ, which produces
a habitat that is vertically constrained to surface waters by low oxygen concentrations, thereby
resulting in increased interactions among prey and predators. This model does not preclude the
possibility that changing food quality associated with the rise of the OMZ, either through deleterious
effects of diatoms on copepods (Ianora et al. 2004) or by low nutrition, might negatively impact
the population. Indeed, a recent study (Vargas et al. 2006b) has shown that available food resources
may strongly affect reproduction and recruitment of zooplankton in the coastal zone of the HCS.
Future perspectives in zooplankton research

Much of the current research devoted to understanding the ecological and biogeochemical role of
zooplankton consumers in the upwelling region off northern Chile is focused on establishing
217
MARTIN THIEL ET AL.
Growth rate
Mortality rate
t Po
en p ula
itm tio
e cr u nd
o dr e ca
p y
pe Population size
Co
Maximum bloom
TEMPERATURE
DIATOMS
OMZ
SpringSummer AutumnWinter
Figure 6 A conceptual model to describe the annual cycle of dominant zooplankton species at the coastal
upwelling off Chile. The model proposes a life cycle in two phases. The first phase (left side) shows a positive
growth of the population (growth rate is stable as well as mortality rate: bottom-up control). The population
reaches a maximum size coinciding with the maximal diatom bloom and then it suddenly collapses. The
second phase (right side) describes the population decay under increased mortality rate of early stages (top-
down control) and a slight decrease of growth rate. The panel below describes the environmental conditions
upon which both phases take place. Temperature has a weak signal, diatoms reach a maximum, but remain
stable, and a key feature is the rise of the oxygen minimum zone (OMZ) coinciding with the population
collapse.
quantitative relationships between species and trophic levels or functional groups (e.g., H.E.
González et al. 2004b). It is expected that trophic relationships may provide insights on the
functional role of zooplankton for recycling C and N in the ocean (Morales 1999, Hidalgo et al.
2005a, Vargas et al. 2007). This should be considered as baseline information for zooplankton
modelling in the region. At population level, modelling approaches have rarely been applied to
zooplankton in this area (Marín 1997). Nevertheless, biogeochemical modelling incorporating zoo-
plankton is an expected issue for coming years. At any level, however, key oceanographic processes
interacting with zooplankton populations need to be identified and understood. Among such pro-
cesses coastal upwelling, OMZ distribution, ENSO variability and changing food quantity and
quality are to be considered crucial.
Fish consumers
The pelagic food webs in the HCS, as in other upwelling systems, feature relatively short trophic
pathways. In general, besides zooplankton (mainly copepods and euphausids) three trophic levels
of consumers can be distinguished: small-size planktivorous fish, larger fish predators and top
predators (Neira et al. 2004). The dominant species among the small-size fish consumers are
anchovy (Engraulis ringens) and Pacific sardine (Sardinops sagax); large predators include the jack
218
mackerel (Trachurus murphyi), hake (Merluccius gayi) and cephalopods (Neira & Arancibia 2004).
Top predators in the HCS are large pelagic fish such as tuna (Thunnus orientalis) and swordfish
(Xiphias gladius), southern sea lions (Otaria flavescens) and seabirds.
Most fish predators in the HCS appear to be non-specialists, which feed opportunistically on
a wide range of different prey items. Sardines and anchovy consume small food particles (mainly
phytoplankton and copepods), with the anchovy able to consume larger food items than sardines
(Balbontín et al. 1979). Anchovy are more specialised on large zooplankton, while sardines consume
a wide range of food items from phytoplankton to small zooplankton (Alheit & Niquen 2004). Jack
mackerel prey on copepods, euphausids, sardines and anchovies and benthic resources (Medina &
Arancibia 2002). Jumbo squids (Dosidicus gigas) feed cannibalistically and on fish, including jack
mackerel and anchovy (Chong et al. 2005). Some of the main fish consumers themselves are prey
to larger predators, for example, swordfish (Ibáñez et al. 2004) and sea lions (Sielfeld 1999). A
generalised scheme of the pelagic food web off central Chile shows the short trophic pathways and
the predominance of relatively few main fish consumers (Figure 7A). While the food spectra of
most pelagic consumers are relatively well known, information about consumption rates, intra- and
interspecific competition or intraguild predation is limited. Similarly, relatively little is known about
prey preferences and feeding strategies of the pelagic fish consumers in the HCS. This knowledge
is particularly important in light of variable oceanographic conditions, which may modify avail-
ability of preferred food items or produce a spatial segregation of predators and prey (e.g., Bakun
2001, Alheit & Niquen 2004).
A review by Cury et al. (2000) suggested that the availability of small pelagic fish in the HCS
off Peru determines the population size of higher trophic levels. Bakun (2001) pointed out that
the interaction between small pelagic fish and their predators is highly dynamic, depending on
multiple environmental (oceanic fronts, climate-driven changes in oceanography), biological (repro-
ductive strategies) and behavioural (schooling behaviour) factors. Small fish consumers may tem-
porarily escape from predation in this dynamic system, and if their reproductive and behavioural
strategies permit them to find a refuge from predation these species may build up and maintain
huge populations (Bakun 2001). Once this dynamic balance is interrupted (e.g., by climatic factors),
regime shifts may occur (Alheit & Niquen 2004).
It has been proposed that the regime shift from a sardine-dominated system to an anchovy-
dominated system (or vice-versa), which is related to long-term variations in oceanographic conditions
(Chavez et al. 2003), may ultimately be mediated by trophic feedbacks (Alheit & Niquen 2004).
During warm periods, the preferred prey items of anchovy (large copepods and euphausids) become
less available (see also Zooplankton consumers, p. 214ff.) while predation pressure on adult anchovies
increases, due to invasions of jack mackerel into coastal waters (Alheit & Niquen 2004). Simulta-
neously, sardines, which are also important predators on anchovy eggs (Alheit 1987), may be favoured
because they have a wide prey spectrum including phytoplankton. The consequences of these regime
shifts, which have been analysed for Peru and northern Chile (Alheit & Niquen 2004), also extend
to central Chile. Future studies need to examine to which degree bottom-up or top-down mechanisms
are involved, and whether EN impacts and fisheries may accelerate these regime shifts.
Both prey and predator behaviour also affects trophic interactions in the HCS. In a recent study,
Bertrand et al. (2006) suggested that the feeding behaviour of jack mackerel is closely linked to
the OMZ and DVM of their prey. These authors discussed that during the day, when prey are hiding
in deeper waters, jack mackerel rest near the upper limit of the OMZ. At night, when prey migrate
into upper oxygenated waters, jack mackerel become active (Figure 7B). If this model is confirmed
in future studies, interannual variation in oceanographic conditions (e.g., the intensity and depth
of the OMZ) may also affect the trophic efficiency of jack mackerel. This hypothetical scenario
underscores the importance of better knowledge of predator–prey behaviour and interactions in
order to better understand the pelagic food webs in the HCS.
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MARTIN THIEL ET AL.
A
pelagic fish predators
sea lions
FISHERY
jack mackerel
cephalopods seabirds
anchovy euphausids
sardines gelatinous
zooplankton
copepods
phytoplankton
B Day Night
100m
jack mackerel
Resting − +
Foraging
200m Hydrologically driven Concentration Trophically driven
Zooplankton
300m
400m Mesopelagic fish
Depth Oxygen
Figure 7 (A) Simplified pelagic food web in the HCS off northern and central Chile; size of boxes indicates
relative proportions; grey arrows show mortality due to fishing. (B) Conceptual model of feeding behaviour
of jack mackerel (Trachurus murphyi) in response to diel vertical migration of prey into the OMZ; position of
jack mackerel shown as grey patches, position of prey organisms shown as continuous grey band. (Modified
after Bertrand et al. 2006)
Interannual fluctuations in fish stocks do not seem to affect the general characteristics of the
food web off central Chile (Neira et al. 2004). However, fisheries appear to have a long-term impact
on the pelagic food web in this area. Fisheries are acting on several trophic levels of the pelagic
food web (Figure 7A). Off central Chile, a decrease in the trophic level of the principal fisheries
resources has been observed during recent decades (Arancibia & Neira 2005). It needs to be taken
into account, however, that populations of some top predators that have experienced severe exploi-
tation in the past (whales) may still be at very low levels, which may be reflected in the structure
of present-day food webs. The interpretation of trophic relationships in the different parts of the
220
HCS (and in other EBCs) requires detailed understanding of the different components of the
respective system (Moloney et al. 2005).
Seabirds and marine mammals

In marine environments seabirds and partially marine mammals are highly visible wide-ranging,
upper trophic-level consumers. Upwelling systems are generally characterised by high production
and relatively short food chains/webs, enabling massive energy transfer to the higher trophic levels
(Cushing 1971, Arntz & Fahrbach 1991). The upwelling system of the Humboldt Current is
particularly well suited for studying the effects of the marine environment on the biology and
ecology of seabirds and marine mammals.
Seabirds
In the HCS there is a rich diversity of seabirds, comprising at least 14 breeding species, 9 of which
are endemic (Table 2). According to available information, along the coast of Chile the most
important breeding colonies are found on islands of northern-central Chile, near upwelling areas.
The grey gull (Larus modestus), which nests inland in the Atacama Desert, travels to the coast on
a daily basis to forage in the upwelling zones off Antofagasta (~23°S). Although proximity to a
feeding ground is relevant in endemic seabirds, it seems that inaccessibility to predators and human
intrusion strongly determines the distribution of breeding populations. Despite some protective
status, most islands in the HCS are subject to human disturbance. In the past, this was caused
mainly by guano harvesting and egg collecting. At present, introduced mammals and unregulated
tourism (Ellenberg et al. 2006) are a major problem.
The majority of the endemic seabirds breed once a year. However, in their northernmost
breeding areas (Peru), species such as the Humboldt penguin (Spheniscus humboldti) nest through-
out the year. Apparently, the two-peak breeding strategy has evolved in response to more favourable
oceanographic and climatic conditions off Peru, and this behaviour is preserved in northern-central
Chile providing additional offspring to those produced during the spring event (Simeone et al.
2002). Furthermore, differences in food availability along its breeding range (~4000 km) might
induce lower breeding performance in southern colonies of this species (Hennicke & Culik 2005).
These birds feed on the main pelagic fish species from the HCS, mainly anchovy and jack mackerel.
The proportion of fish stocks taken by endemic seabirds is not well known, but most likely does
not exceed 10%. Incidental mortality of endemic seabirds due to fisheries is mainly caused by gill
nets (Simeone et al. 1999, see also Majluf et al. 2002).
The HCS is visited regularly by a number of migrant species. Among the Procelariiformes, white-
chinned petrels (Procellaria aequinoctialis), Buller’s albatrosses (Thalassarche bulleri), Antarctic
prions (Pachyptila desolata) and Juan-Fernández petrels (Pterodroma externa) are the most abundant
species during austral summer (Weichler et al. 2004). There is evidence that the HCS is also frequented
by other remarkable visitors such as the Chatham, wandering and royal albatrosses (Thalassarche
eremita, Diomedea exulans, and D. epomophora, respectively). Apparently the presence of these
species at such a distance from their colonies is related to the food abundance in the HCS, which in
summer also attracts species like the black-browed albatross (Thalassarche melanophris) from south-
ern islands (56°S) (Arata & Xavier 2003) and during winter species like the white-chinned petrels
from South Georgia (Phillips et al. 2006). The abundances of some of these visitors during austral
summer may reach 2.5–5 birds km−2, but at present it is not known which proportion of the entire
population of these visitors may at times use the HCS as a feeding ground. The large albatrosses
mainly feed on cephalopods and pelagic fishes (resulting in deadly interactions with long-line fisheries),
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MARTIN THIEL ET AL.
Table 2 Breeding abundance and conservation status of seabirds (number of pairs) and mammals
(individuals) on the coast of northern-central Chile. The geographic divisions correspond approxi-
mately to the limit of political division of Chile. Data for cetaceans (ψ) correspond to systematic
counts during oceanographic trips conducted every January (1999, 2002, 2003 and 2004) surveying
an area of ~420 km2. The other data for cetaceans are mainly from sightings made from the land.
Species 18–21°59′S 22–26°59′S 27–29°59′S 30–32°59′S 33–34°S Total CS*
Seabirds
Order Sphenisciformes
Humboldt penguin Spheniscus 8120 810 8930 V
humboldti φ
Magellanic penguin Spheniscus 5 5 NT
magellanicus
Order Procellariiformes
Peruvian diving-petrel Pelecanoides 4170 4170 E
garnotii φ
Wedge rumped storm petrel 9 9 LC
Oceanodroma tethys
Order Pelecaniformes
Peruvian booby Sula variegata φ 15,560 15,560 LC
Peruvian pelican Pelecanus 400 1000 1400 LC
thagus φ
Neotropic cormorant Hypoleucos 200 50 55 305 —
brasiliensis
Guanay cormorant Leucocarbo 12 150 162 NT
bougainvillii φ
Red legged cormorant Stictocarbo 122 85 60 23 150 440 NT
gaimardi φ
Order Charadriiformes
Band tailed gull Larus belcheri φ 1 1 LC
Grey gull Larus modestus φ 10,000 10,000 LC
Kelp gull Larus dominicanus 2700 100 450 3250 LC
Inca tern Larosterna inca φ 10 10 NT
Peruvian tern Sterna lorata φ 2499 E
Marine mammals
Order Carnivora
Family Otariidae
South american fur seal 655 1205 1860 LR
Arctocephalus australis
South american sea lion Otaria 13,797 6132 129 1598 21,656 LR
flavescens
Family Mustelidae
Marine otter Lontra felina 1.5 1.0–2.5 E
(ind./km of coast)
Order Cetacea (numbers come from sightings)

Family Balaenidae
Southern right whale Eubalaena 21 21 LR
australis
222
Table 2 (continued) Breeding abundance and conservation status of seabirds (number of pairs)
and mammals (individuals) on the coast of northern-central Chile. The geographic divisions correspond
approximately to the limit of political division of Chile. Data for cetaceans (ψ) correspond to systematic
counts during oceanographic trips conducted every January (1999, 2002, 2003 and 2004) surveying
an area of ~420 km2. The other data for cetaceans are mainly from sightings made from the land.
Species 18–21°59′S 22–26°59′S 27–29°59′S 30–32°59′S 33–34°S Total CS*
Family Balaenopteridae
Humpback whale Megaptera 1 17 ψ 18 V
novaeangliae
Balaenoptera sp. 25 9ψ 34
Fin whale Balaenoptera physalus 63 ψ 63 E
Minke whale Balaenoptera 1 4ψ 5 LR
acutorostrata
Blue whale Balaenoptera 1ψ 1 E
musculus
Family Kogiidae
Dwarf sperm whale Kogia simus 3ψ 3
Family Physeteridae
Sperm whale Physeter 1 1 19 ψ 20 V
macrocephalus
Family Ziphiidae
Cuvier’s beaked whale Ziphius 5 5 —
cavirostris
Family Delphinidae
Bottlenose dolphin Tursiops 301 ψ 301 DD
truncatus
Short-finned pilot w. n.d. LR
Globicephala macrorhynchus
Long finned pilot whale 101 101 LR
Globicephala melas
Globicephala sp. 11 ψ 11
Killed whale Orcinus orca 11 ψ 11 LR
False killer whale Pseudorca 5ψ 5 LR
crassidens
Common dolphin Delphinus 200 200 LR
delphis
Southern rightwhale dolphin 450 450 DD
Lissodelphis peronii
Risso’s dolphin Grampus griseus 30 ψ 30 DD
Dusky dolphin Lagenorhynchus 456 1404 ψ 1860 DD
obscurus
Family Phocoenidae
Harbour porpoise Phocoena 72 10 ψ 82 DD
spinipinnis
Notes: *CS: Conservation status was extracted from IUCN (2006). E: Endangered, V: vulnerable, NT: near threatened,
LC: least concern, DD: data deficient, LR: lower risk. φ Endemic species of the Humboldt Current System. Data from
Guerra et al. 1987, 1988, Sielfeld et al. 1997, van Waerebeek et al. 1998, Capella et al. 1999, Sielfeld & Castilla 1999,
Rendell et al. 2004, Simeone et al. 2003, Frere et al. 2004, Jiménez 2005, Bernal et al. 2006, and G. Luna-Jorquera
unpublished data.
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MARTIN THIEL ET AL.
while the smaller petrels consume mainly planktonic crustaceans (e.g., Euphausia mucronata).
Although available information shows that incidental mortality in the artisanal Patagonian toothfish
(Dissostichus eleginoides) fisheries in southern Chile is low (0.047 birds per 1000 hooks; C.A. Moreno
et al. 2006), incidental mortality of petrels and albatrosses in industrial long-line fisheries has not yet
been assessed for northern Chile, where most of the Chilean swordfish vessels are based.
Marine mammals
Species of mammals listed in Table 2 are those that mainly occur over the continental shelf in the
HCS. Although there are no endemic species, the total species richness reaches at least 22 species,
most of them being cetaceans. The number of species and abundance of individual sightings in
Table 2 suggest that the upwelling regions between 18°S and 30°S are important feeding stations
on the migration routes of whales (see also Rendell et al. 2004).
For the sea otter (Lontra felina), available information shows that this species feeds mainly in
areas dominated by macroalgae (Ebensperger & Castilla 1992, Ebensperger & Botto-Mahan 1997,
Villegas 2002), where refuges on land are available (Sielfeld & Castilla 1999). The diet of this
species is principally composed of coastal fishes, crabs and seabirds (Sielfeld & Castilla 1999,
Mattern et al. 2002) and foraging success is higher at wave-protected sites than at exposed sites
(Villegas et al. 2007).
The biology and ecology of otariids at both sea and land has been investigated in Peru (Majluf
et al. 2002) and Ecuador (e.g., Dellinger & Trillmich 1999), but studies from northern-central Chile
mostly focus on population size or demography (e.g., Guerra et al. 1987). Interaction between sea
lions (Otaria flavescens) and fisheries is an important issue in Chile (Hückstädt & Krautz 2003).
Their traditional foraging behaviour (search and pursuit of prey) has changed to one of ‘sit and
wait’ for food captured by fishing vessels (Hückstädt & Antezana 2003), a strategy that appears to
be exploited by killer whales (Orcinus orca) preying on sea lions near fishing vessels (Hückstädt
& Antezana 2004). While it is well known that sea lions destroy nets and consume pelagic fish,
the extent of this problem has not yet been systematically examined in northern Chile.
Although there is a lack of information, it appears that there exists an inverse relationship between
the breeding distribution of sea lions and endemic seabirds in northern-central Chile (Table 2). Sea
lions are most abundant between 18°S and 26°S, while seabirds seem to be more abundant between
27°S and 33°S. The reasons for this spatial separation are not well known, but one possibility might
be that sea lions are common in northern Chile where they can successfully breed on small rocky
outcrops, while seabirds may be most abundant between 27°S and 33°S, where they form huge
breeding colonies on undisturbed islands. Additionally, maternal care might impose higher restric-
tions (duration and distance of foraging trips) on female sea lions (e.g., Trillmich 1990) than on
female seabirds, which share parental care with their male partners and thus can afford longer absence
from the nest and farther foraging trips during breeding. The apparent lower productivity of pelagic
fish between 27°S and 33°S (see also Pelagic fisheries and fisheries management 1980–2005, p. 288ff.)
might affect female sea lions (which usually forage close to the breeding sites; e.g., Trillmich 1986)
to a higher degree than seabirds (which may travel >30 km from breeding colonies during individual
foraging trips, as observed in Humboldt penguins; Luna-Jorquera & Culik 1999). Future studies
should examine the causes for the inverse abundance patterns of sea lions and seabirds along the
coast of northern Chile.
Effects of ENSO on seabirds

In the upwelling area of Coquimbo (Region IV), the species composition of seabirds at sea fluctuates
among different sites, and distribution patterns are linked to variation in SST and chlorophyll
224
concentration. Dense swarms of pelagic fishes attract high numbers of piscivorous birds, which is
the most important factor to explain their distribution patterns. Usually, only 3% of all birds recorded
in a given area are seen directly attending fishing vessels (Weichler et al. 2004). Life-history theory
predicts that seabirds will respond to reduction in food availability by changing their behaviour
and/or breeding effort, thus buffering adult survival. Data of Peruvian boobies (Sula variegata) are
in agreement with this prediction (G. Luna-Jorquera unpublished data). In January 2002, about
3000 breeding pairs of this species were registered in a colony off Coquimbo, but 1 month later
there were only 10 pairs, coinciding with a positive SST anomaly (Figure 8). Counts of birds at
sea conducted in ~420 km2 around the colony showed that the numbers of boobies were lower than
in a ‘normal’ year, indicating that boobies left the area probably due to a reduction in food
availability. This caused both a reduction in the total numbers of all seabirds foraging in feeding
flocks and a simultaneous increase of seabirds attending fishing vessels. During cooler LN years,
large concentrations of seabirds were seen in feeding flocks and few birds followed fishing boats
(Figure 8). The apparent change in food availability is also reflected in the high number of breeding
pairs of boobies during LN years (Figure 8).
Endemic seabirds have evolved in the upwelling area of the HCS, where historically their
populations are subjected to large fluctuations, ENSO being an important selective force for the
evolution of their breeding biology and life-history patterns. During EN, some species abruptly
leave their eggs and young to undertake eruptive movements away from the usual breeding and
feeding sites in apparent search for food; total or partial breeding failure then often occurs (Simeone
et al. 2002). Seabirds are generally characterised by longevity, high age of first breeding, slow
reproductive rate and intense offspring care and are thus typical K-strategists. However, it has been
hypothesised that, compared with related species, Peruvian boobies and Guanay cormorants
(Leucocarbo bougainvilli) have larger clutches, may attempt to breed more than once within 1 yr
and reach sexual maturity at an unusually early age (Luna-Jorquera et al. 2003). This enables them
to build up populations rapidly in the years after EN events because even young, unexperienced
adults are able to raise large broods because the food supply per bird is much greater than it would
be in a population in equilibrium with the environment (Furness & Monaghan 1987). However,
since the establishment of the anchovy fishery, the dynamics of the seabird populations have changed
(Duffy et al. 1984). Instead of rapidly increasing populations by raising large broods at least once
per year, endemic seabirds failed to respond to the reduced competition brought about by their
reduction in numbers. It seems that the anchovy fishery has taken up the superabundance of food,
which in the past permitted the seabirds from the HCS to cope with the recurring crashes induced
by oceanographic perturbations (Jahncke et al. 2004). In fact, data available for Peruvian seabird
colonies (Tovar & Cabrera 1985) show that the EN 1957–1958 caused a mortality of 39% of the
estimated populations of 28 million adult birds. Mortality increased to 57% (of 6.54 million adult
birds) due to EN 1972–1973, after 1970 when anchovy landings had reached 12 million t (metric
tons). The extreme EN 1982–1983 caused a decrease from 6 million birds to only 0.3 million birds
in Peru. After that, the populations showed an increase in numbers, reaching ~6 million birds, but
the EN 1997–1998 reduced it again to less than 0.4 million birds. In 2000, the size of the Peruvian
seabird population was estimated to be 1.93 million adult birds, showing a slight recovery (IMARPE
2006). No comparable datasets are available for the coast of northern-central Chile, where only
infrequent surveys of breeding colonies have been conducted so far (e.g., Simeone et al. 2003).
Food resources or breeding sites?

Seabirds and sea lions feed on fish and population crashes during EN years have been explained
with the disappearance of fish stocks. Which proportion of the fish stocks do seabirds and seals
consume? And how are they affected by variable food supply? Bioenergetic modelling provides
225
MARTIN THIEL ET AL.
temperature
2
anomaly °C
Sea surface
1
0
−1
−2
−3
10000
Nesting pairs
February
1000
January
Numbers
100
10
n.d.
Boobies at sea
Numbers
2000
1000
n.d. n.d.
1999 2000 2001 2002 2003 2004
Total birds
Numbers
4000
2000
n.d. n.d.
0
Percent of all seabirds at sea
30 Percent in feeding flocks

20
10
n.d. n.d. 0
30 Percent attending
fishing vessels
20
10
n.d. n.d. 0 0
1999 2000 2001 2002 2003 2004
Figure 8 Abundance of breeding and foraging seabirds in the vicinity of Coquimbo, Chile (30°S), and the
relationship with sea-surface temperature (SST) anomalies in the southeast Pacific. Upper panel shows SST
anomalies during the study period; middle panel shows nesting pairs of Peruvian boobies in a breeding colony
on Pájaros Island (29°35′S), and boobies counted during censuses at sea in the Upwelling system of Coquimbo,
between 29°08′ and 30°11′ (see Weichler et al. 2004 for more details); lower panel shows total number of all
seabirds counted in the study area in January of each year, and the percentage of birds foraging in feeding
flocks or attending fishing vessels in search of food.
useful quantitative assessment of the impact of marine vertebrates on fish stocks and may serve to
predict changes in fishery practices or effects of fish stock sizes on seabird and seal numbers.
Simple predictions would be that species with specialised feeding methods and a high dependence
on specific diets, which have been reduced in availability (due to fisheries or oceanographic
226
conditions), would be most likely to decline in numbers. In order to develop the models to test
these predictions, a long-term research programme is necessary to assess (1) breeding populations,
(2) feeding ecology, (3) reproductive biology and (4) energy budgets, under different environmental
conditions. In particular, it appears to be important to examine the breeding biology of seabirds.
There is an indication that populations of seabirds in northern and central Chile may be limited by
the availability of undisturbed breeding islands (see Schlatter 1984, 1987, Schlatter & Simeone
1999). While additional research is necessary, present knowledge already identifies the protection
of undisturbed breeding sites as one of the highest priorities for seabird conservation in the HCS
of northern and central Chile.
Sandy beaches
Sandy beaches are a common feature of the Chilean coast between 18°S and 41°S (Figure 9), where
they offer breeding habitats and important food resources for migrating shorebirds. The extent and
distribution of sandy beaches offers unique opportunities for studying the factors driving the
population dynamics of invertebrate consumers along the coast of northern and central Chile.
Beaches are found along exposed shorelines, sheltered bays and coastal islands and their total extent
and average length show a clear latitudinal trend, increasing toward southern-central Chile
(Figure 9). Sandy beaches of the coast of Chile show a general zonation pattern of three zones that
differ both in their physical characteristics and biological communities (Figure 10) (Jaramillo 1987,
Average distance Average length Total extent

between beaches (km) of beaches (km) of beaches (km)
6 4 2 0 6 4 2 0 300 200 100 0
−18°
−20°
−22°
−24°
−26° Southern latitude
−28°
−30°
−32°
−34°
−36°
−38°
−40°
−42°
A B C
Figure 9 (A) Average distance (mean ± standard error) between beaches, (B) average length of individual
beaches, and (C) total extent of sandy beaches within each sector; individual sectors correspond to the coastline
between two subsequent degrees of latitude (information extracted from GOOGLE-EARTH, a total of 810
sandy beaches were measured between 18°S and 42°S).
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MARTIN THIEL ET AL.
Winter Summer
9
2
5
1 −20°
8 10
−25°
Southern latitude
6 3 −30°
4
−35°
−40°
Low Mid High Low Mid High
Food supply
Habitat harshness
Figure 10 Zonation scheme of sandy beach community along the coast of Chile during winter and summer;
shaded background areas represent intertidal gradients in food supply (FS, dotted area) and habitat harshness
(HH, grey area), showing both latitudinal and seasonal variation in these physical factors; size of boxes with
animal sketches represents proportional abundances of the respective species; note the latitudinal difference
in species composition in the upper intertidal zone; species shown are 1, Ocypode gaudichaudii; 2, Phalerisida
maculata; 3, Orchestoidea tuberculata; 4, Tylos spinulosus; 5, Excirolana braziliensis; 6, E. hirsuticauda; 7,
E. monodi; 8, Emerita analoga; 9, Nephtys impressa; 10, Mesodesma donacium. (Figure inspired by a drawing
from Jaramillo 1987.)
McLachlan & Jaramillo 1995, Jaramillo et al. 2001). The lower and middle shore have similar
species composition (i.e., Emerita analoga, Mesodesma donacium, Nephtys impressa, Excirolana
braziliensis, E. hirsuticauda and E. monodi) along the continental coast of Chile, while the upper
shore has species-composition differences between northern, central and southern Chile (Jaramillo
1987) (Figure 10). The upper zone, extending from the drift line up to the dunes, is a very dry
environment, occasionally moistened by sea mist. Inhabitants of this zone, such as tenebrionid
beetles (Phalerisida maculata), talitrid amphipods (Orchestoidea tuberculata) and oniscoid isopods
(Tylos spinulosus) feed on plant and animal remains washed up on the beach (Jaramillo 1987). In
the northern zone (18–25°S) and off Peru, the amphipod and isopod species are replaced by the
ghost crab Ocypode gaudichaudii (Quijón et al. 2001), which together with Phalerisida maculata
inhabits the upper shore (Figure 10). To which degree species replacement is due to physical factors
or to species interactions is not well known at present. However, the fact that these faunal changes
in the supralittoral zone are accompanied by a gradient in burrowing depth and proximity to the
drift line of the main organisms suggests that physical factors (in particular desiccation risk) may
play an important role. Guppy (1906) expressed that “the beaches are of dry loose sand in which
the hand fails to find on scooping below the surface that refreshing coolness which is the character
of beaches in all latitudes where the land is vegetated”.
228
Within the beach, there are differences not only in the interspecific but also in the intraspecific
zonation patterns, which may be accompanied by behavioural differences. In particular, locomotor
activity and position on the beach varies depending on the developmental stage and on the time of
the day. For example, juvenile Orchestoidea tuberculata and Phalerisida maculata are most active
during the day, while adults show highest activities at night (Jaramillo et al. 1980, 2000). This has
been suggested as a strategy to avoid intraspecific competition for food and additionally the risk
of intraspecific predation. Furthermore, Orchestoidea tuberculata shows an aggregated distribution
on kelp patches stranded in the supralittoral zone (Duarte et al. 2004), which is related mainly to
feeding behaviours. For Emerita analoga and Mesodesma donacium, similar ontogenetic differences
in activity or habitat use have been reported. Juvenile Emerita analoga typically occur in highest
abundances in the low intertidal zone slightly above the adults (Contreras et al. 2000). Similarly,
juvenile Mesodesma donacium inhabit the lowest intertidal zone of the beach (in the swash zone),
while adults live in permanently water-covered sediments of the subtidal zone (mainly in the surf
zone) (Jaramillo 1994).
Most sandy beach studies have centred on the effect of physical factors, in particular wave
exposure, grain size and tides, on the community structure. Species richness, biomass and abundance
decrease from dissipative to reflective beaches (Jaramillo 1994, McLachlan & Jaramillo 1995).
Biological interactions are more intense on dissipative beaches than on reflective beaches, where
the population dynamics are mainly controlled by physical factors (Defeo & McLachlan 2005).
Strong competition between mole crabs Emerita and surf clams Donax has been suggested as the
cause for the aggregated distribution of the surf clams (Leber 1982), and negative interactions
between Emerita analoga and Mesodesma donacium were observed along the Chilean coast (Dugan
et al. 2004), but aggregations may also be a response to sediment characteristics (i.e., grain size).
On the other hand, organisms inhabiting the upper shore (i.e., talitrid amphipods) are not associated
to one particular beach type (Defeo & McLachlan 2005), which suggests that their populations are
not influenced by beach morphodynamics but rather by other physical factors or food availability.
To which degree resource or interference competition (or direct predation) is responsible for the
latitudinal changes in the species composition of this zone remains to be explored.
The species that inhabit sandy beaches have diverse reproductive strategies, which can be related
to the habitat characteristics. For example, most of the species of the upper and middle shore (i.e.,
Orchestoidea tuberculata, Tylos spinulosus and Excirolana spp.) have direct development, which
might suggest that the connectivity between populations is low. In northern-central Chile (29°S)
the reproductive peak of Excirolana hirsuticauda is in spring–summer (Contreras & Jaramillo
2003), and it is suggested that the other species from the upper/middle shore follow a similar
pattern. The lower intertidal and subtidal zone is inhabited by organisms with indirect development
(Emerita analoga, Mesodesma donacium, Nephtys impressa), which have planktonic larvae possibly
permitting higher connectivity between local populations. For Emerita analoga, distinct recruitment
peaks (see also Arntz et al. 1987 for Peru) have been reported for autumn, spring or early summer
in northern (22°S) and southern-central Chile (39°S), but ovigerous females are usually found
throughout the year (Contreras et al. 1999, Contreras et al. 2000). In northern-central Chile (30°S)
the surf clam Mesodesma donacium has two spawning peaks during the year (usually during the
early summer and autumn), but females with mature gonads have been observed throughout most
of the year (Alarcón & Navea 1992, Stotz et al. 1999). These observations suggest continuous
reproductive activity, but spawning or successful recruitment seems to occur only infrequently,
possibly depending on particular environmental factors (triggering gamete release) and oceano-
graphic conditions (affecting larval survival and supply).
There is also a high temporal variability of faunal abundance throughout the year. For example,
Orchestoidea tuberculata reach highest abundances during the winter months, both in central (30°S)
and southern Chile (40°S) (Sánchez et al. 1982, McLachlan & Jaramillo 1995). Furthermore, during
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the winter, these amphipods live higher up on the beach, mainly to avoid being washed out by
higher wave activity. During summer, the abundances of O. tuberculata decrease and they are found
closer to the flotsam (Sánchez et al. 1982, McLachlan & Jaramillo 1995), possibly to avoid
desiccation risk in the supralittoral zone or to get close to their food. The cirolanid isopods
(Excirolana spp.) have similar abundances in summer and winter, but they shift their position in
the intertidal zone, moving higher during the winter months (Sánchez et al. 1982). On the other
hand, Emerita analoga in the northern (22°S) and Mesodesma donacium in northern-central Chile
(30°S) reach highest abundances in summer (Sánchez et al. 1982, Contreras et al. 2000). At present,
the reasons for the apparent inverse population dynamics of species from the upper versus those
from the lower intertidal zone are speculative, and seasonally varying offspring (larval) survival or
food supply could be invoked. One very interesting species from the supralittoral zone, Tylos
spinulosus, which has a restricted distribution in northern-central Chile (17°–30°S) (Schmalfuss &
Vergara 2000), is little known aside from a few data on its population density (Sánchez et al. 1982,
Jaramillo et al. 2003), and it appears a promising enterprise to examine its population dynamics.
The vicinity to upwelling centres plays an important role in the succession and structure of
hard-bottom communities (Broitman et al. 2001, Narváez et al. 2006), but there is no clear indication
that the macrofauna composition of sandy beaches is influenced by their proximity to upwelling
areas (Jaramillo et al. 1998). Contreras et al. (2000) concluded that growth rates of Emerita analoga
from a beach near the upwelling centre of Mejillones (22°S) were within values reported for other
areas, suggesting only limited or no direct effects of upwelling on sandy beach inhabitants. Com-
munity dynamics of sandy beaches may be influenced by upwelling to a lesser degree than those
pertaining to hard-bottom communities. For example, higher nutrient availability near upwelling
areas positively influences growth rates of seaweeds on hard bottoms (Camus & Andrade 1999,
Wieters 2005) and thereby the community succession (Nielsen & Navarrete 2004), which clearly
is of no importance on exposed sandy beaches where algae are usually imported from neighbouring
(or distant) hard-bottom habitats. Furthermore, the interplay between upwelling and subsequent
relaxation events strongly affects the recruitment of hard-bottom organisms with planktonic larvae
(Narváez et al. 2006), but seems to be of minor importance on sandy beaches, where the most
common organisms feature direct development.
The effect of ENSO has been intensively studied in hard-bottom environments, but its role in
the dynamics of sandy beach communities is not well known (Arntz et al. 1987). EN events may
have deleterious effects on the organisms from the lower intertidal zone of sandy beaches (e.g.,
Mesodesma donacium; see also Artisanal benthic fisheries, p. 278ff.). On the other hand, it is known
that EN provokes mass mortalities of seaweeds and animals, many of which eventually will strand
on sandy beaches (Arntz 1986). This high supply of OM may represent an important food source
for scavenging animals of the supralittoral zone of sandy beaches (e.g., Orchestoidea tuberculata).
In this way, it can be suggested that intertidal organisms are distinctly affected by EN: species from
the lower shore (suspension feeders with planktonic larvae) may be negatively affected by EN,
while those from the supralittoral zone (scavenging animals with direct development) might benefit
from the higher food supply and more benign climate (lower desiccation risk) during EN.
Subtidal soft-bottom communities

Zonation patterns
Recent studies suggest that the bathymetric distribution of subtidal benthic communities off the
Chilean margin seems to be controlled mainly by bottom water oxygen conditions and sediment
organic loading. OMZs are significant mid-water features in the eastern Pacific Ocean (Wyrtki
1973, Kamykowski & Zentara 1990) that strongly influence the distribution and diversity of
230
planktonic and benthic marine communities. Where OMZs intercept the continental margin (bottom-
water dissolved oxygen < 0.5 ml L−1), strong gradients are formed in both bottom-water oxygen
concentration and OM input (Levin et al. 1991, Levin et al. 2000). These gradients influence the
biogeochemical properties of sediments (Cowie et al. 1999) and the distribution and diversity of
bacteria, meio-, macro- and megabenthic organisms (Sanders 1969, Mullins et al. 1985, Wishner
et al. 1990, Tyson & Pearson 1991). Off Chile, oxygen-deficient waters are in general associated
with the ESSW, which partially covers the continental shelf and upper bathyal area. The intensity
and vertical extent of the OMZ suggest a latitudinal gradient, the effect disappearing at about 41°S
(Brandhorst 1971). Off northern Chile, sediments affected by the OMZ extend from a few tens of
metres below the surface to 300–400 m water depth. Between Huasco and Valparaíso (~28–32°S),
the OMZ seems to intercept the sea floor deeper than 100 m (D. Lancellotti & W. Stotz unpublished
data), and due to the narrowness of the shelf and steepness of the slope, there are zones that probably
are not severely affected. This is also corroborated by the presence of a fauna atypical for oxygen-
deficient areas (e.g., diverse species of gastropods) and the absence of bacterial mats (Lancellotti
& Stotz 2004). The shelf widens southward and when upwelling prevails, during spring–summer,
the OMZ again can be found only a few metres from the surface even in southern-central Chile
(~36°S), within Concepción Bay (Ahumada et al. 1983), and extending down to 200–300 m (see
also Arntz et al. 2006). However, the OMZ intensity here is probably the result of many local
factors (e.g., the high PP that leads to high remineralisation rates in the water column and sea floor,
consuming oxygen and generating sulphidic conditions within the sediment) (P. Muñoz et al.
2004b). In this way, it is probably possible to visualise the OMZ impinged sea floor, from northern
to central Chile, as a wedge-shaped band, getting narrower southward, but with two foci of most
intense oxygen-deficient conditions, one off northern Chile and the other at the shelf off Concepción.
The continuity between these two foci may be interrupted by better-oxygenated sediments, at
comparable depths, off central Chile. Reports of low bivalve abundances between 80 and 120 m
depth in Valparaíso Bay (31°S) are suggestive of OMZ effects (Ramorino 1968), but additional
data are required to resolve the intensity and extent of the OMZ and its effect on benthic commu-
nities between 25°S and 35°S.
Considering the general effect of the OMZ on benthic communities, and based on the limited
amount of biological sampling available at that time, Gallardo (1963) proposed the existence of
basically three main benthic zones for the local eukaryotic communities: (1) an upper sublittoral
zone, up to 50 m depth, with favourable conditions for the development of ‘normal’ benthic
communities, (2) a lower sublittoral zone, from 50 to 300–400 m (varying with latitude and
coinciding with the extent of the OMZ), in which only those organisms highly adapted to cope
with oxygen deficiency and high organic loadings are able to thrive (basically small polychaetes,
oligochaetes, nematodes and a few molluscs), and (3) a bathyal area, associated mainly with Antarctic
Intermediate Waters, with a diverse and rich fauna (dominated by annelids, crustaceans, molluscs
and echinoderms) that benefits from enhanced oxygen and good quality and quantity of sediment
OM. How this general pattern differs in southern areas (>41°S) where the OMZ dissipates is still
poorly known.
One of the most distinguishing features of benthic shelf communities within OMZ-impinged
sediments is the presence of extensive mats of the filamentous, sulphide-oxidising bacteria Thio-
ploca and Beggiatoa (Gallardo 1963, 1977, Schulz et al. 2000, Arntz et al. 2006). These bacteria
are the most conspicuous component of the benthos also in the central and southern Peruvian shelf
(Rosenberg et al. 1983). Bacterial biomasses of up to 1 kg m−2 wet wt have been reported from
shelf sediments off Iquique (~21°S) (Gallardo 1963) and off Concepción (~37°S) (Gallardo 1977)
at depths between 50 and 100 m. On the other hand, within the OMZ eukaryotes are in general
small-size forms, like meiofauna, calcareous foraminiferans and nematodes (Gooday et al. 2000,
Neira et al. 2001, Levin 2003). Very high densities, on the order of 10,000 individuals (ind.) 10 cm–2,
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MARTIN THIEL ET AL.
of meiobenthic organisms, mostly nematodes, have been recorded on the shelf off Concepción
(Neira et al. 2001, Sellanes et al. 2003). OMZ macrofaunal assemblages thus have low diversity
and are typically composed of organisms with morphological and metabolic adaptations and feeding
strategies suited to these conditions. A good example is the locally abundant polychaete Parapri-
onospio pinnata, an interface feeder organism, which is also able to switch to suspension feeding
(Gutiérrez et al. 2000), with a complete set of enzymes for anaerobic metabolism (González &
Quiñones 2002), and also bearing a set of highly branched and complex gills. Paraprionospio
pinnata comprises about half of the ~14,000 macrofauna individuals m−2 reported from off Con-
cepción within the OMZ (Palma et al. 2005).
Below the OMZ, the total abundance of macrobenthos decreases while biomass and diversity
increase in parallel with increases in oxygen and decreases in organic carbon (Levin 2003). In the
bathyal area off Chile, important crustacean assemblages, mainly of commercially valuable galat-
heid decapods (e.g., Pleuroncodes monodon and Cervimunida johni), thrive at the slope within and
below the lower boundary of the OMZ. Below the OMZ, groups that are favoured by increased
oxygen concentrations are decapod crustaceans, gastropods and at greater depths echinoderms
(mainly ophiuroids, asteroids and irregular echinoids). The large tubiculous onuphid polychaete
Hyalinoecia sp. (tubes 10–20 cm in length) is particularly abundant at mid-slope depths off
Concepción and Chiloé (J. Sellanes personal observations). Scleractinian corals and many species
of gorgonarians are common on the slope off Chiloé, below 500 m water depth.
Finally, though the three-layer scheme for benthic zonation is the general rule, another type of
chemosynthetic community, methane seep areas, has been reported from off Concepción at
700–1400 m water depth (Stuardo & Valdovinos 1988, Sellanes et al. 2004, Sellanes & Krylova 2005).
In these reducing systems, C is fixed locally by both free-living and symbiotic chemosynthetic bacteria.
Seepage areas thus provide a suitable environment for the development of singular communities
consisting of sulphide oxidising bacteria (e.g., Beggiatoa), highly endemic endosymbiont-bearing
clams (e.g., Vesicomyidae, Lucinidae, Thyasiridae and Solemyidae) and tubeworms (e.g., Lamelli-
brachia sp.). In addition, non-chemosymbiotic megafauna (e.g., crustaceans, gastropods, cephalo-
pods, fish) are massively attracted to these deep-sea hot spots of biological activity. The attraction
of these areas is due to both the abundance of locally produced organic material and the presence
of authigenic carbonate reefs (generated by microbial and chemical processes), which are avidly
colonised by a diverse benthic fauna (J. Sellanes unpublished data).
Latitudinal and bathymetric patterns

In general, for the available data of macrofaunal species richness on the Chilean shelf, no clear
latitudinal pattern emerges. Published reports of species numbers range from 18 to 56 species at
northern Chile (~23°S, 15–65 m water depth) (Jaramillo et al. 1998), 42 to 85 species at Huasco
(~28°S, 20–50 m water depth) (Lancellotti & Stotz 2004), 10 to 41 species at Concepción (~36°S,
15–65 m water depth) (Carrasco et al. 1988), and 24 to 34 species further south (~45°S)
(D. Lancellotti & W. Stotz unpublished data). Highest abundances reported are for the shelf off
Concepción (36,290–38,590 ind. m−2; Carrasco et al. 1988) and biomass values of almost 900 g
wet wt m−2 are reported for south of Chiloé (D. Lancellotti & W. Stotz unpublished data).
A crude comparison among the few available deeper studies (>100 m water depth) suggests
higher macrofaunal standing stocks and abundances also at the shelf off central and southern areas,
both for macro- (body size 0.3–2 cm) and megafauna (body size >2 cm), compared with northern
Chile (Figure 11). Palma et al. (2005) reported macrofaunal biomass values for three transects
covering a depth range from about 100 m to 2000 m at Antofagasta, Concepción and Chiloé (about
22°S, 36°S and 42°S, respectively). Though biomass trends with depth, beyond the OMZ, are in
general unimodal at all transects, with higher values at intermediate depths and lowest within the
232
Megafaunal abundance
Macrofaunal abundance
100000 70
Antofagasta (∼22°S) Macrofaunal biomass
Megafaunal species number 60
10000
Macrofaunal species number
Macrofaunal biomass (g wet weight m−2), Megafaunal and macrofaunal species number
Megafaunal abundance (Log10 ind 1000 m−2), Macrofaunal abundance (Log10 ind m−2) 50
1000 40
100 30
20
10
10
1 0
98 142 300 520 690 1350 1800
10000 70
Concepción (∼36°S)
60
10000
50
1000 40
100 30
20
10
10
1 0
120 365 535 800 1290 2050
10000 70
Chiloé (∼42°S)
60
10000
50
1000 40
100 30
20
10
10
1 0
160 300 480 900 1250 1960
Depth (m)
Figure 11 Macro- and megafauna depth-related patterns for three transects (22°S, 36°S and 42°S) across the
shelf and upper bathyal zone of the Chilean margin. Data for macrofaunal abundance, biomass and species
number from Palma et al. (2005) and for megafaunal abundance and species number from E. Quiroga
unpublished data).
OMZ and beyond 1350 m, average biomasses are lower off Antofagasta. Maximum values for this
transect (6.9 g wet wt m−2) are reported at 518 m water depth, while values about an order of
magnitude higher (60.7 g wet wt m−2) are reported off Concepción at 784 m depth. For southern
Chile (~42°S) intermediate values (39.2 g wet wt m−2) are reported for a station located at 1250 m
depth. This also indicates a deepening of macrofaunal biomass maxima with latitude (Figure 11).
For the megafauna observed at the same three transects, though biomass values are not available,
abundances in general exhibited a similar pattern to that previously explained for macrofaunal
233
MARTIN THIEL ET AL.
biomass (Palma et al. 2005). On average, pooling the data for the three transects, abundance (~500
ind. m−2) and species number (~25) peaks were located between 1000 and 1500 m depth.
Temporal patterns of variability in shelf communities

As explained in previous sections of this review, the coastal zone off northern and central Chile is
strongly influenced by seasonal wind-driven upwelling, giving rise to one of the areas with the
highest PP rates known worldwide (Fossing et al. 1995, Daneri et al. 2000). Since water column
oxygen conditions and sediment organic loadings fluctuate at both intra- (i.e., seasonal) and
interannual scales (i.e., ENSO cycle), some degree of coupling with the dynamics of benthic
communities is expected (Tomicic 1985, Arntz & Fahrbach 1991). This has been demonstrated for
southern-central Chile (~36°S), where seasonal and interannual changes in upwelling intensity can
lead to changes in bottom-water dissolved oxygen concentration, in the amount of OM reaching
the bottom (Gutiérrez et al. 2000), in the quality and lability of deposited OM (Neira et al. 2001,
Sellanes & Neira 2006) and in the sediment nitrogen fluxes (P. Muñoz et al. 2004b). During the
last strong EN event (1997–1998), important insights were gained by examining the effects of
changing environmental conditions on local bacterial, meiofaunal (Neira et al. 2001, Sellanes et al.
2003) and macrofaunal (Gutiérrez et al. 2000) communities off central Chile. The largest biomasses
of the bacterial component have been observed after several years of upwelling-favourable condi-
tions, which are associated with cold LN phases of the ENSO (V.A. Gallardo et al. unpublished
data), while the bacterial biomass is effectively depressed during warm EN phases. A decreasing
trend in macrofaunal density, as well as the presence of deeper-burrowing infauna, evolved toward
the end of EN 1997–1998, mainly due to the decrease of the polychaete Paraprionospio pinnata
(Gutiérrez et al. 2000). It appears that more oxygenated bottom waters and oxidised sediment during
EN caused P. pinnata to fail in its summer recruitment. In addition, it is probable that increased
competition and predation by other species have contributed to its decline. Indeed, it has been
reported that during EN, many subtropical predators invade the coastal areas (Arntz et al. 1991),
negatively affecting the surface-feeding polychaetes (Tarazona et al. 1996). In central Chile,
P. pinnata recovered its numerical dominance only in summer 2003, i.e., 5 yr after the end of EN.
Severe hypoxic and sulphidic conditions that developed during summer 2003 probably eliminated
or precluded possible competitors and/or predators, triggering the explosive increase of the
P. pinnata population during this period (Sellanes et al. 2007).
In northern Chile (20–30°S) few datasets extending over at least 12 months are available from
shallow benthic communities (20–80 m). For the time period 1990–1995, relatively high abundances
of polychaetes have been reported from several stations in northern Chile (23°50′S) at water depths
of 50–60 m (Carrasco 1997). This author remarked on the absence of a clear seasonal signal in
abundance changes of the main polychaete species, and he suggested that the observed variations
rather reflected long-term patterns. At a long-term monitoring station near 28°S, abundances of
polychaetes were high in 1995 (Figure 12), comparable to those found by Carrasco (1997). High
abundance, biomass and species diversity at 28°S were associated with the warm period 1993–1995
and followed by a strong decline in 1996, coincident with LN conditions, which continued during
the EN 1997–1998 (Lancellotti 2002, D. Lancellotti & W. Stotz unpublished data). Gradual disap-
pearance of spionids, as observed in Huasco in 1996 during LN (Figure 12), was also observed
during the same time period in Iquique (20°S) between 9 and 30 m depth (Quiroga et al. 1999).
In northern-central Chile, during EN events, increased wave activity and freshwater runoff are
frequent, in contrast to calmer periods recorded during LN events. Turbulence and runoff, in a zone
where rains and strong storms are uncommon, probably oxygenate the water column, resuspend
OM and/or provide terrigenous material, thus favouring reproduction and settlement of mac-
robenthic species living below the zone of direct wave and sediment deposition impact (>20 m
234
6000 60
Upper sublittoral zone
5000 50
28°S
4000 40
3000 30
2000 20
Abundance (ind m−2)
Species richness
1000 10
0 0
6000 60
Lower sublittoral zone (OMZ)
5000 50
36°S
4000 40
3000 30
2000 20
1000 10
0 0
93 94 95 96 97 98 99 2000 01 02 03 04 05
Years
Figure 12 Temporal variations in the abundance of soft-bottom macrofauna in the upper sublittoral zone
(northern Chile, 28°S) and the lower sublittoral zone (central Chile, 36°S); average abundance and species
richness (S) are given; data from the upper littoral zone are taken by Smith-MacIntyre grab (1995–1996) or
with sediment cores by divers (1997–2005) (D.A. Lancellotti & W. Stotz unpublished data); data from the
lower sublittoral zone are taken with a multicorer, and several samples from each year were pooled, thus not
allowing intra-annual variation to be seen (J. Sellanes unpublished data).
water depths). Effects of EN (and other) events on the temporal variability of benthic soft-bottom
communities at present are difficult to evaluate because very few long-term datasets from benthic
habitats are available from the HCS along the Chilean coast. It is herein suggested that long-term
monitoring programmes should be implemented, sampling on a seasonal or bimontly basis, fol-
lowing examples in Peru (Tarazona et al. 2003, Arntz et al. 2006, Peña et al. 2006) and the Northern
Hemisphere (Frid et al. 1996, Kroencke et al. 1998, Salen-Picard et al. 2002).
Intertidal and subtidal hard-bottom communities

Hard bottoms along the coast of northern-central Chile are generally restricted to a narrow fringe
extending from the intertidal zone to shallow sublittoral waters. The rock substratum is composed
of rock of volcanic, granitic or sedimentary origin (Fariña et al. in press). The extensive rocky
shores between 18°S and 40°S are mostly exposed to strong wave action, and they are only
interrupted by short stretches of sandy beaches, which increase in extent toward the south (see
Sandy beaches, p. 227, Figure 9), thereby leading to a wider separation of neighbouring hard-
bottom environments. Communities on intertidal and subtidal hard bottoms are dominated by
macroalgae and suspension-feeding animals that form large patches (occasionally extending over
to neighbouring soft bottoms) or belt-like stretches (running parallel to the shore at a certain tidal
level). Most intertidal and subtidal hard bottoms are covered by one or a few dominating habitat-
forming organisms. Patches may persist for many years at a given location (Durán & Castilla 1989,
Fernández et al. 2000, Vega et al. 2005), and they offer abundant microhabitat and food for asso-
ciated organisms (Moreno & Jara 1984, Vásquez et al. 1984, Núñez & Vásquez 1987, Buschmann
1990, Vásquez 1993a, López & Stotz 1997, Sepúlveda et al. 2003a,b). Here these habitat-forming
235
MARTIN THIEL ET AL.
species, their spatial and geographic extent, their temporal dynamics and their role as ecosystem
engineers (EEs) will be described. Further, it will be briefly discussed how the spatiotemporal
distribution of these EEs may influence local biodiversity, population dynamics and trophic inter-
actions in hard-bottom communities along the HCS.
Habitat-forming species on hard bottoms

Large kelps (Lessonia nigrescens, L. trabeculata, Macrocystis integrifolia, M. pyrifera and Durvil-
laea antarctica), which grow abundantly in the low intertidal and shallow subtidal zone of the HCS,
have compact and complex holdfasts that offer abundant and diverse microhabitats on the rock
substratum itself. Their stipes and blades reach lengths of 2.5 m (Lessonia trabeculata) up to 30 m
(Macrocystis pyrifera), and they have an important effect on local hydrodynamics because they act
as wave breakers and slow down currents (Graham 2004). Smaller macroalgae with shorther thalli
of 5–50 cm (such as Halopteris funicularis, Glossophora kunthii, Asparagopsis armata, Corallina
officinalis and Gelidium chilense) form dense carpets (turfs) that offer primary and secondary
microhabitats because they act as sediment traps retaining sand and shell fragments between their
thalli and stolons (López & Stotz 1997, Kelaher & Castilla 2005).
A diverse group of suspension-feeding EEs on hard bottoms include polychaetes (Phragmato-
poma moerchi), barnacles (Austromegabalanus psittacus), bivalves (Perumytilus purpuratus, Semi-
mytilus algosus, Choromytilus chorus and Aulacomya ater), and ascidians (Pyura chilensis and
P. praeputialis). Their matrices reach heights of 2–40 cm, offering abundant space between living
individuals (Cerda & Castilla 2001) or in remaining tubes or shells of dead individuals. Matrices
of these suspension feeders may also retain considerable amounts of sediments (Prado & Castilla
2006), thereby providing secondary substratum for associated organisms.
Habitat-forming species compete among themselves for space on hard-bottom substrata. Several
studies indicate that mussels are competitively superior over barnacles (Navarrete & Castilla 1990,
2003, Tokeshi & Romero 1995a) and can also overgrow turf algae (Wieters 2005), ascidians may
outcompete mussels (Castilla et al. 2004a) or barnacles (Valdivia et al. 2005), and large kelp may
recruit in and then overgrow turf algae (Camus 1994a). Superior competitors themselves may be
suppressed by predators (e.g., mussels and ascidians by gastropod and seastar predators; Paine et al.
1985, Castilla 1999, Castilla et al. 2004b) and grazers (Vásquez & Buschmann 1997, Buschmann
et al. 2004a). Humans, acting as top predators by removing intermediate consumers, also strongly
influence the structure of hard-bottom communities in northern and central Chile (Moreno et al.
1986, Castilla 1999). Furthermore, recruitment and growth of habitat-forming species are controlled
by a variety of processes (e.g., upwelling) that drive larval and food supply (Navarrete et al. 2002,
Nielsen & Navarrete 2004, Wieters 2005). Following disturbances and detachment, open space on
hard bottoms is quickly recolonised, starting with ephemeral algae, which subsequently are replaced
by large and long-lived turf or kelp algae and suspension feeders (Durán & Castilla 1989; Valdivia
et al. 2005).
Spatial and temporal dynamics of ecosystem engineers

The geographic range of most macroalgae and suspension-feeding EEs extends throughout northern-
central Chile (into Peru), but not all of them have a continuous latitudinal distribution (Table 3).
All EEs from hard bottoms have pelagic dispersal stages, but in the case of the macroalgae the
planktonic phase is of short duration (minutes to hours).
Kelps of the genera Lessonia and Macrocystis extend from southern Chile to north of 18°S,
but EN events may provoke large-scale extinctions in northern Chile (see also Kelp forests,
236
Table 3 Main ecosystem engineer species from intertidal and subtidal hard bottoms along the Humboldt Current System of northern and central
Chile
Zonation Wave- Height of Distance between Patch persistence
Ecosystem engineer (m) exp. patches (m) Size of patches (m2) Latitudinal extent (°) patches (km) (years)
<1
1 to 10
10 to 1000
> 1000
18,19
20,21
22,23
24,25
26,27
28,29
30,31
32,33
34,35
36,37
38,39
<1
1 to 10
10 to 1000
> 1000
<1
1 to 10
>10
Lessonia trabeculata 0–30 E,S,P 2.5 x x x x x x

Lessonia nigrescens 0 E,S 6 x x x x x x
Macrocystis integrifolia 0–15 S,P 10 x x x x x x
Macrocystis pyrifera 0–30 S,P 30 x x x x x
Durvillaea antarctica 0 E 15 x x x x x x
Glossophora kunthii 0–8 E,S,P 0.25 x x x x x x
Halopteris funicularis 0–8 E,S,P 0.1 x x x x x x
Asparagopsis armata 0–8 S,P 0.12 x x x x x x
Corallina officinalis 0–8 E,S,P 0.15 x x x x x x
Gelidium chilense 0–2 E,S,P 0.06 x x x x x
237
Phragmatopoma moerchi 0 E 0.2 x x x x
Perumytilus purpuratus 0 E 0.05 x x x x x x x x
Semimytilus algosus 0 E 0.1 x x x x x x
Aulacomya ater 10–20 E 0.2 x x x x x x x
Austromegabalanus psittacus 0–20 E,S 0.2 x x x x x
Pyura chilensis 1–20 E,S,P 0.2 x x x x x x
Pyura praeputialis 0–10 E,S,P 0.6 x x x x x x
Notes: temporally extinct

common and widespread
present, but patchy
rare
absent
Information obtained from Hoffmann & Santelices 1997, Santelices 1989, Castilla et al. 2000, Zagal & Hermosilla 2001, Sepúlveda et al. 2003a,b and Hernández et al. 2001.
MARTIN THIEL ET AL.
p. 240ff.). Durvillaea antarctica does not extend further north than 32°S. Patches or belts formed
in the low intertidal zone by Lessonia nigrescens and Durvillaea antarctica generally have extents
of several square metres up to >100 m2. Subtidal kelp forests of Lessonia trabeculata and Macro-
cystis spp. may extend over >1000 m2, comprising some of the largest habitat patches formed by
EEs. Distances between neighbouring patches are small in the case of Lessonia spp. and Durvillaea
antarctica but individual forests of Macrocystis spp. can be separated by several hundred kilometres
(see also Kelp forests, p. 240ff.). Persistence of patches over time may be favoured by recruitment
of new sporophytes into existing kelp patches (Santelices & Ojeda 1984).
Turf algae generally form smaller patches, with individual patches rarely exceeding an area of
a few square metres. Corallina officinalis and Gelidium chilense (and other turf algae, e.g., Mon-
temaria horridula, Rhodymenia skottsbergii) form long-lived patches in the intertidal zone (López
& Stotz 1997, Vásquez & Vega 2004a, Wieters 2005), and distances between neighbouring patches
are relatively small (Table 3). Glossophora kunthii, Halopteris funicularis, Asparagopsis armata
(and others, including Corallina officinalis) occur mainly on shallow subtidal hard bottoms where
they form patches of several square metres and, while thalli may disappear during the winter, the
stolons persist over several years (Vásquez et al. 2001a). Patches may extend their size or renew
thalli via asexual proliferation.
The polychaete Phragmatopoma moerchi forms patches of several square metres in extent in
the low intertidal and shallow subtidal zone in areas with a high supply of sand and shell fragments
(Sepúlveda et al. 2003b). These patches persist over several years, but disappear if renewal is
reduced, either due to low larval supply or high postsettlement mortality (Zamorano et al. 1995).
The barnacle Austromegabalanus psittacus forms aggregations in the low intertidal and shallow
subtidal zone; patches generally are small, rarely exceeding more than a few square metres in area.
This species occurs all along the coast of northern and central Chile and little is known about the
temporal dynamics of individual patches. Bivalves form extensive patches of a few square metres
up to >1000 m2 in area in the mid-intertidal (Perumytilus purpuratus), low intertidal (Semimytilus
algosus) and subtidal zones (Choromytilus chorus, Aulacomya ater). In the absence of predators
patches can persist over many years (Durán & Castilla 1989), facilitated by regular recruitment
into adult patches (Alvarado & Castilla 1996). Most bivalve species have a wide latitudinal
distribution, but Perumytilus purpuratus and Aulacomya ater are almost entirely absent over an
extensive area in northern Chile between 23°S and 32°S (Fernández et al. 2000, personal observa-
tions), which appears to be mainly due to limited larval supply in that region (for Perumytilus
purpuratus see Navarrete et al. 2005). The ascidian Pyura chilensis occurs in small patches in the
shallow subtidal zone (e.g., Vásquez & Vega 2004b), while the congener P. praeputialis forms
extensive belts in the low intertidal zone (Table 3). Patches of P. chilensis persist over many years at
the same location (personal observations), but little is known about the population dynamics within
patches. In P. praeputialis, recruitment may be most successful in the vicinity of adults (Clarke
et al. 1999), thereby favouring the long-term persistence of patches. While P. chilensis has a wide
geographic distribution, P. praeputialis is restricted to a small range of 70 km along the Bay of
Antofagasta (23°S) in northern Chile (Castilla et al. 2000).
Macrofauna associated with ecosystem engineers on hard bottoms

A wide diversity of organisms is associated with habitat-forming species on hard bottoms of
northern and central Chile. Highest species richness is found in the kelp holdfasts, intermediate
numbers of associated species are reported from ascidian and bivalve reefs, and turf algae harbour
fewest species of associated macrofauna (Figure 13A). This relationship appears to be related to
238
140 120
120 100
Species richness
100
80
80
60
60
40
40
20 20
0 Corallina officinalis 0
Gelidium chilense
Lessonia trabeculata
Pyrua praeputialis
Lessonia nigrescens
Phragmatopoma moerchi
Perumytilus purpuratus
Perumytilus purpuratus
Durvillaea antarctica
Halopteris funicularis
Macrocystis integrifolia
Macrocystis integrifolia
1 to 10 10 to 1000 >1000
Patch size (m2)
B
Kelps Turf algae Suspension feeders

A
Figure 13 (A) Species richness of macroinvertebrates associated with habitat-forming macroalgae or suspen-
sion feeders from intertidal and subtidal hard bottoms of the northern and central coast of Chile; for reasons
of comparability only studies that reported at least seven phyla of associated macrofauna were considered.
(B) Average species richness in biotic habitats of different patch sizes; information obtained from López &
Stotz 1997, Gelcich 1999, Godoy 2000, Thiel & Vásquez 2000, Cáceres 2001, Cerda & Castilla 2001,
Hernández et al. 2001, Vásquez et al. 2001b, Thiel & Ullrich 2002, Sepúlveda et al. 2003a,b, Prado & Castilla 2006.
the fact that kelp beds and ascidian and bivalve reefs have a comparatively large spatial extent
while patches of turf algae rarely cover more than a few square metres (Figure 13B). A positive
relationship between patch size and number of associated species has been revealed for most habitat-
forming species (Vásquez & Santelices 1984, Villouta & Santelices 1984, Thiel & Vásquez 2000,
Hernández et al. 2001, Sepúlveda et al. 2003a,b).
Several macrofauna species have been reported from a variety of different biotic habitats. Of
251 species identified from biotic substrata (see references in Figure 13), 11.6% have been found
in all three types of main biotic habitats (kelps, turf algae and suspension feeder reefs), 23.5% have
been found in two types and 64.9% are only reported from one type of habitat. It must be emphasised
that so far no single study has compared the associated fauna among the three main types of EEs,
and there is little indication that there are habitat specialists that only occur in one type of biotic
substratum. For example, Hernández et al. (2001) emphasise that several of the polychaetes found
in patches of the barnacle Austromegabalanus psittacus also occur in other habitats. Similarly,
Sepúlveda et al. (2003b) mention that many species from surrounding habitats associate with the
reef-building polychaetes Phragmatopoma moerchi. They also emphasise that these biotic substrata
may serve as recruitment habitat for some organisms. Similar observations led López & Stotz
(1997), who found juvenile stages of many crustaceans and molluscs in Corallina officinalis, to
239
MARTIN THIEL ET AL.
speak of a ‘transitory fauna’ in biotic substrata (see also Vásquez & Santelices 1984, Cerda &
Castilla 2001). EEs may thus favour many mobile organisms that temporarily find shelter in these
habitats (e.g., Vásquez et al. 2001b). In this context, Castilla et al. (2004b) reported that the intertidal
ascidian Pyura praeputialis facilitates the extension of mobile macrofauna from the subtidal into
the mid-intertidal zone, thereby enhancing local species richness.
The main functional groups of the organisms associated with biotic habitats are suspension
feeders (32.4% of all species), grazers (25.2%) and predators (23.4%) (H. Bastias & M. Thiel
unpublished data). Vásquez et al. (2001b) found very similar proportions of functional groups both
in kelp holdfasts and on the surrounding hard bottoms. By offering structural protection, EEs are
considered to mediate species interactions and buffer the effect of physical stress, often favouring
suspension feeders (Wieters 2005, Valdivia & Thiel 2006). While the role of EEs in sustaining and
promoting local biodiversity on intertidal and subtidal hard bottoms has been elucidated in numerous
studies during the past decades (Vásquez & Santelices 1984, Villouta & Santelices 1984, Cerda &
Castilla 2001, Sepúlveda et al. 2003a,b, Castilla et al. 2004b, Prado & Castilla 2006), relatively
little is known about their trophic role on exposed rocky shores of northern-central Chile. Several
studies have underlined the role of kelp forests as contributors of algal biomass to neighbouring
habitats (Rodríguez 2003) and as feeding grounds for fish predators that consume understory algae
and kelps (Angel & Ojeda 2001) or associated fauna (Núñez & Vásquez 1987, Palma & Ojeda
2002). Fish consumers are known to play an important role in kelp food webs of northern-central
Chile (Angel & Ojeda 2001, Fariña et al. in press) but little is known about the food webs in other
EEs. While most studies acknowledge the importance of EEs as habitat for associated organisms,
their trophic efficiency (uptake of nutrients and suspended matter, release of dissolved and partic-
ulate organic matter) and the role of associated macrofauna in the tropho-dynamics of communities
on intertidal and subtidal hard bottoms have not been thoroughly studied (see also Graham et al.
2007). The high biomass and diverse assemblage of associated consumers suggest that EEs are
energetic power plants that concentrate and convert food resources in a similar way to kelp, seagrass
or suspension feeder reefs in other parts of the world (e.g., Asmus & Asmus 1991, Lemmens et al.
1996, Wild et al. 2004).
Kelp forests
Giant kelp dominate shallow, subtidal rocky-bottom areas in temperate and cold seas down to a
depth of ~40 m (Dayton et al. 1984, Harrold & Pearse 1987, Vásquez 1992, Graham et al. 2007).
Kelp distribution from south Peru to central Chile is as follows: (1) intertidal rocky areas are
dominated by Lessonia nigrescens, which forms belts along exposed coasts; (2) rocky subtidal
environments are dominated by Lessonia trabeculata until 40 m in depth; (3) Macrocystis integri-
folia forms shallow kelp beds from the intertidal zone to depths of about 15 m. In southern-central
Chile, these species are gradually replaced by Durvillaea antarctica, which dominates the intertidal
zone in wave-exposed areas (Hoffmann & Santelices 1997), and in subtidal areas by Macrocystis
pyrifera, which occurs in both wave-exposed and –protected habitats (Buschmann et al. 2006a).
While the two species from the genus Lessonia have an almost-continuous distribution along the
entire Chilean continental coast, Macrocystis integrifolia has a fragmented distribution, forming
patchy populations in northern Chile. In this zone Lessonia trabeculata and Macrocystis integrifolia
coexist, but mixed kelp populations have segregated patterns of bathymetric distribution, M. integri-
folia being more abundant in shallow areas (Vega et al. 2005). Local populations may vary from
hundreds of metres to hundreds of kilometres in extent. The observed distribution patterns are the
result of complex life-history strategies that interact with environmental factors such as spatial and
temporal variation in water movement, nutrient availability and temperature (Buschmann et al.
2004b, V. Muñoz et al. 2004, Vega et al. 2005).
240
The kelp forest community

Kelp communities are highly productive (Dayton 1985), and they harbour a high diversity and
abundance of invertebrates and fishes. Kelps, especially their holdfasts, constitute feeding areas,
refuges against predation and bottom currents, spawning, settlement areas and nursery sites
(Vásquez & Santelices 1984, Vásquez et al. 2001c, Vásquez & Vega 2005). Below the kelp canopy
a wide diversity of turf algae exists, including several Corallinales, Asparagopsis armata, Halopteris
paniculata and Gelidium spp.; several species of barnacles and other sessile invertebrates (Pyura
chilensis, Phragmatopoma moerchi, Aulacomya ater) are also part of the associated species shel-
tered by the kelp canopy (Vásquez et al. 2001b,c, Vásquez & Vega 2004a). In contrast to the
Northern Hemisphere, no large predators have been reported for southeastern Pacific kelp beds
(Graham et al. 2007). Instead, invertebrate predators such as the muricid snail Concholepas conc-
holepas, seastars (Meyenaster gelatinosus, Stichaster striatus, Heliaster helianthus and Luidia
magellanica), and intermediate-size coastal fishes (Cheilodactylus variegatus, Semicossyphus mac-
ulatus and Pinguipes chilensis) dominate the predator guild in kelp forests from northern-central
Chile (Vásquez 1993b, Vásquez et al. 1998, 2006). These predators feed on a diverse guild of
herbivores, including sea urchins (Tetrapygus niger and Loxechinus albus), gastropods (Tegula spp.
and Fissurella spp.), as well as fishes (Aplodactylus punctatus, Girella laevifrons and Kyphosus
analogus) (e.g., Medina et al. 2004). These herbivore species graze on kelp and associated algae,
regulating their abundance and distribution (Vásquez & Buschmann 1997, Vega et al. 2005, Vásquez
et al. 2006). Marine mammals widely distributed in the coastal zone of the HCS, such as sea lions
Otaria flavescens and sea otters Lontra felina, also use kelp beds as feeding areas.
Population dynamics and spatial distribution of kelps in northern-central Chile

The kelp species from northern and central Chile belong to the Laminariales, which have a complex
life cycle with two morphologically different stages: one conspicuous stage, recognisable as kelp
that produces spores (the sporophytes), and a microscopic stage comprising independent female
and male plants (the gametophytes) that lead a hidden life in the benthos. Sporophytes are the
product of gametic reproduction, which is triggered by environmental factors (temperature, irradi-
ance, photoperiod, and nutrient concentrations).
The sporophytes themselves are reproductive year-round, but peak spore release has been
observed during winter. Since spore survival in Laminariales is short, the dispersal range of kelps
is generally assumed to be quite reduced (Graham et al. 2007); if spores do not settle within a
relatively short period they die (Santelices 1990a). However, spores may survive in the guts of
different herbivores (Santelices & Correa 1985, Santelices & Payá 1989) or as filaments in darkness
(Santelices et al. 2002). Furthermore, fertile floating plants may act as spore carriers and thereby
contribute to dispersal (Macaya et al. 2005).
Juvenile sporophytes of Lessonia recruit onto hard-bottom substrata during late winter–spring,
and in the field are capable of producing spores after 6–8 months (Santelices & Ojeda 1984; see
review by Edding et al. 1994). Interference by adult plants inhibits the intertidal recruitment of
juvenile L. nigrescens in exposed habitats. Nevertheless, water movements produce whiplash effects
(sensu Dayton et al. 1984) that gives protection against grazers, thereby promoting successful
recruitment of sporophytes (Santelices & Ojeda 1984). In subtidal habitats abundance of grazers,
currents and reproductive behaviour of two species of elasmobranchs (Schroederichthys chilensis
and Psammobatis scobina) affect Lessonia trabeculata populations. Grazing modifies algal mor-
phology, producing two morphotypes: shrub-like and tree-like morphs. Water movement affects
these differentially and generates higher mortality on tree-like morphs (Vásquez 1992). Short
distances between plants (or high densities) reduce the access of grazers to the holdfasts. The
241
MARTIN THIEL ET AL.
whiplash effect of fronds and stipes pushes herbivores away from the plants, reducing grazing
pressure. On the other hand, spawning of egg cases of elasmobranchs on L. trabeculata ties the
stipes together, thereby reducing the whiplash effect and thus permitting grazers to approach kelp
plants. Additionally, this ‘tie effect’ modifies plant shape toward the tree-like morph, and plants
are more easily dislodged by water movement (Vásquez 1992).
Longevity of kelps from northern Chile in the field is not well known since they do not show
any evident age-related structure. Nevertheless, individual Lessonia plants can survive in the field
for as long as 5 yr (J.A. Vásquez personal observations), and Macrocystis integrifolia has been
reported as a perennial species in northern Chile (Buschmann et al. 2004b). Several factors generate
significant biomass loss in the field: grazing pressure, wave impact, and spore release, which takes
place mainly during summer (Santelices & Ojeda 1984, Edding et al. 1994).
Lessonia and Macrocystis populations in northern-central Chile grow throughout the year but
exhibit growth peaks during spring–summer (Buschmann et al. 2004b, Tala et al. 2004). Growth
patterns are modified by wave impact, quantity and quality of light, water temperature and nutrient
concentration (Buschmann et al. 2004b, Vega 2005). Local factors such as intraspecific interactions
(Santelices & Ojeda 1984), herbivory (Vásquez & Buschmann 1997, Vásquez et al. 2006) and
coastal upwelling events (González et al. 1998, Vásquez et al. 1998) can modify seasonal patterns
of abundance and distribution (see also Graham et al. 2007). Large-scale phenomena such as ENSO
produce interannual variability in abundance and could eventually generate local extinctions, as
observed after the EN events of 1982–1983 and 1997–1998 (Soto 1985, Tomicic 1985, Vega 2005,
Vásquez et al. 2006). Major impacts of EN were observed in kelp beds from lower latitudes
(18–21°S). For example, a kelp bed occupying an area of ~40 ha at 18°S during the 1970s (IFOP
1977) disappeared as a consequence of EN 1982–1983 (Soto 1985) and has not recovered since.
Similarly, during EN 1997–1998, the density of adult sporophytes on subtidal hard bottoms at 21°S
decreased rapidly and linearly with increasing positive thermal anomalies (Figure 14). Six months
later the site remained completely devoid of adult sporophytes, and no recolonisation occurred in
the subtidal zone during the study period. In areas south of 23°S positive thermal anomalies
registered during EN 1997–1998 had only limited effects on kelp beds (Figure 14). As a result, the
spatiotemporal abundance patterns of M. integrifolia sporophytes in northern-central Chile is highly
variable (Figure 14).
Kelp conservation and human activities

Many of the diverse kelp-associated species have significant socioeconomic importance for human
populations along the coast in north-central Chile and have been subject to harvesting by local
human communities since pre-Columbian times (Jerardino et al. 1992, Vásquez et al. 1996). Spatial
and temporal dynamics of kelp beds are significantly affected by anthropogenic impacts produced
by both intense harvesting and severe pollution with organic as well as mining waste (Faugeron
et al. 2005; Vásquez & Vega 2005). Lessonia nigrescens, L. trabeculata and Macrocystis integrifolia
are commercially exploited between 18°S and 32°S. These species account for >95% of landings
of macroalgae and basically are used for alginic acid extraction. Until 2002, collected biomass in
dry weight (dry wt) amounted to ~200,000 t, almost exclusively based on stranded kelps, resulting
from natural mortality of plants with holdfasts that are weakened by grazing and then detached by
strong bottom currents and waves. Since 2003, however, due to international needs for raw (dry)
materials and also due to increasing demands for fresh algae (to sustain aquaculture of herbivorous
invertebrates in northern Chile), harvesting of natural kelp increased to ~300,000 t dry wt per year.
This has led to the recent implementation of new administrative rules in order to mitigate the impact
on natural kelp populations. Regulations aim at the establishment of a sustainable kelp fishery,
applying the following strategies: (1) harvest management (Vásquez 1995, 1999, 2006), (2) stock
242
Thermal anomalies (°C)

sporophytes (ind m−2)
Subtidal kelp
Abundance of adult
5 Camarones (18°S) 5
4 3
Intertidal kelp
3 1
2 −1
5 1 −3
San Marcos (21°S)
4 0 −5
1996 1997 1998 1999 2000
3 1
2 −1
1 −3
0 −5
5 5
Constitución (23°S)
4 3
3 1
2 −1
1 −3
0 −5
5 5
Abundance of adult sporophytes (ind m−2)
Playa Blanca (28°S)

4 3
3 1
2 −1
1 −3
0 −5
5
Thermal anomalies (°C)
5
Los Choros (29°S)
4 3
3 1
2 −1
1 −3
0 −5
5 5
San Lorenzo (30°S)
4 3
3 1
2 −1
1 −3
0 −5
5 5
Los Vilos (32°S) 3
4
3 1
2 −1
1 −3
0 −5
1996 1997 1998 1999 2000
Figure 14 Temporal variation (between 1996 and 2000) of abundances of adult sporophytes of M. integrifolia
(●) and thermal anomalies estimated in situ (line) over a latitudinal gradient in northern Chile. Note: At San
Marcos, an intertidal kelp population appeared after the El Niño event (▫), while the subtidal kelp bed did
not recover. At Camarones (top) no sporophytes were observed during the study period. (Modified from Vega
2005)
243
MARTIN THIEL ET AL.
enhancement (Vásquez & Tala 1995), (3) cultivation (Edding & Tala 2003, Westermeier et al. 2006,
Gutierrez et al. 2006), and (4) conservation programmes including Marine Protected Areas (MPAs;
CONAMA 2006). Considering the high variability of kelp populations in northern Chile, the limited
dispersal capability of Lessonia species, and in particular the patchy distribution of beds of Mac-
rocystis integrifolia, sustainable exploitation of natural kelp forests requires integrated management
plans with continuous monitoring of standing stocks.
Export and import processes within the HCS

Kelp forests, as other EEs, strongly influence trophic fluxes in benthic environments (e.g., Graham
et al. 2007). In the HCS, some of the most important trophic connections occur in the vertical
direction, such as dissolved nutrients released from sediments into the water column, upwelling of
nutrient-rich waters from subsurface layers to the sea surface, or POM sinking from the euphotic
zone toward deeper water layers and finally to the sea floor. In addition, there exist numerous types
of horizontal transfer of particulate or dissolved components between the marine and the terrestrial
realm, between the benthic and the pelagic environment, or between benthic habitats. In the
following section the importance, intensity and frequency of these exchange processes in the HCS
of northern-central Chile are considered, with a focus on coastal habitats.
Exchange between realms

Exchange between the marine and the terrestrial environment occurs in both directions. Flux of
materials toward the sea is via rivers, which (due to limited freshwater flow) is usually only of
minor importance between 18°S and 30°S. In some areas in northern-central Chile, dissolved and
solid components from human activities are continuously supplied to the marine environment,
impacting local intertidal and subtidal communities (Vásquez et al. 1999, Gutiérrez et al. 2000,
Lancellotti & Stotz 2004). During certain time periods (EN events or summer rains in the Andes
highlands), river flow increases dramatically, transporting mainly sediments but also large quantities
of terrestrial vegetation to nearshore coastal waters (Vásquez et al. 2001a). Shallow subtidal habitats
along the coast of northern-central Chile are infrequently impacted by these mud flows (Miranda
2001, Vásquez et al. 2001a, Lancellotti & Stotz 2004), and it is considered likely that these impacts
occur in parallel over a large geographic range.
In the reverse direction, several natural exchange mechanisms are relevant in the HCS, namely,
energy transfer by seabirds and marine mammals from offshore waters to coastal habitats, which
occurs in a highly concentrated manner on breeding or roosting sites. Additionally, some terrestrial
vertebrates (rodents, lizards, songbirds) from coastal environments forage along the drift line or in
the intertidal zone (Navarrete & Castilla 1993, Fariña et al. 2003a, Sabat et al. 2003). Most of these
organisms maintain relatively stable territories along the coastline, and thus material transfer from
the intertidal to the upper supralittoral zone is dispersed in space, but relatively continuous in time.
The same process has been reported from coastal habitats in California, where populations of insects,
spiders, lizards, rodents and coyotes are mainly maintained and modulated by the food subsidy
from the marine environment (Polis & Hurd 1995, 1996, Polis et al. 1997).
Under certain conditions, marine resources are also transported toward the shore without the
aid of biotic agents (invertebrate or vertebrate consumers defecating on land). Mortality of seabirds
and marine mammals during EN events results in large numbers of animal carcasses accumulating
on local beaches (e.g., Guppy 1906, Arntz & Fahrbach 1991). Similarly, during storm events, algae
or benthic invertebrates are detached and cast onto the shore (González et al. 2001). These food
bounties attract large numbers of terrestrial vertebrate scavengers but since supply is highly infrequent
(e.g., Moore 2002), no quantitative estimates of material transfer during these events are available.
244
Fisheries also contribute to the transfer of OM from the marine toward the terrestrial realm.
This not only includes direct (extraction) but also indirect forms of transfer, such as by scavenging
seabirds around fishing vessels at sea (Weichler et al. 2004) or in fishing ports (Ludynia et al. 2005).
Populations of kelp gulls (Larus dominicanus) near main population centres in northern-central
Chile depend to a large extent on these human-derived food sources, and they then distribute remains
in terrestrial environments (Ludynia et al. 2005).
Exchange between environments

There is a wide range of exchange processes between the pelagic and the benthic environments.
This includes, for example, supply of POM from the water column to soft bottoms where micro-
and macroorganisms remineralise this POM, returning dissolved materials to the water column
(Graf 1989, Marcus & Boero 1998, Dunton et al. 2005). Suspension feeders are important agents,
which aid in transfer of suspended material (e.g., phytoplankton and kelp detritus) from the water
column to the benthic system (Wolff & Alarcón 1993). In some of the bays of northern-central
Chile dense stocks of natural or culture beds may significantly affect these fluxes (Uribe & Blanco
2001, Avendaño & Cantillánez 2005). The intensity and direction of transfer can be affected by
regional discontinuities in the oceanographic conditions (e.g., distance from upwelling areas), which
influence the transport and flux of POM and nutrients (Graco et al. 2006). These processes are also
exposed to large-scale temporal variations in oceanographic conditions (e.g., ENSO cycles) (Farías
et al. 2004, P. Muñoz et al. 2004b). Fisheries in small fishing ports contribute POM in the form of
fish remains to soft bottoms (Sahli 2006).
On hard bottoms, macroalgae and suspension feeders take up nutrients and suspended POM
from the water column, returning algal remains, repackaged faeces or dissolved excretions to the
water column. Most large kelps are continuously shedding senescent parts (e.g., Tala & Edding
2005). These authors estimated that annual export of shed plant detritus from a kelp forest of
Lessonia trabeculata may amount to 18 kg wet wt m−2. What proportion of this detritus remains
suspended in the water column or sinks immediately to the bottom is not known at present. Kelp
productivity shows some seasonal variation, but kelp detritus is supplied throughout the year, at
least in northern-central Chile (Tala & Edding 2005). This suspended kelp detritus may also sustain
the large proportion of suspension-feeding organisms on intertidal and subtidal hard bottoms (see
also Intertidal and subtidal hard-bottom communities, p. 235ff. and Bustamante & Branch 1996).
Little is known about the role of DOC released by kelp forests. It enhances bacterial populations
(Delille et al. 1997) and contributes to foam lines at the sea surface, which are thought to play a
role in propagule dispersal and survival (Meneses 1993, Shanks et al. 2003a). Foam lines are
frequently observed along the coast of northern-central Chile.
Fish consumers also have an important role in energy transfer from benthic toward pelagic
environments (Angel & Ojeda 2001). This transfer includes all feeding guilds of fishes from
herbivores to omnivores and carnivores (Angel & Ojeda 2001). Studies of trophic coupling between
hard bottoms and the water column have mainly focused on kelp forests and little is known about
these trophic interactions in other hard-bottom communities (see also Intertidal and subtidal hard-
bottom communities, p. 235ff.).
Exchange between benthic habitats

Exchange between neighbouring communities (NCs) occurs throughout the shallow subtidal zone.
The form of materials exchanged between NCs and the direction of transport can be highly variable.
Algal detritus exported from kelp forests contributes an important food source for animal communities
on intertidal hard bottoms (Bustamante et al. 1995, Bustamante & Branch 1996, Rodríguez-Graña
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MARTIN THIEL ET AL.
Table 4 Different types of material transfer between communities within the HCS of northern
Chile. Distances of transport increase with increasing length of line, intensity of transfer increases
with increasing size, and frequency augments with increasing number of dots.
Material type Agent Distance Intensity Frequency Reference
Between Realms TERRESTRIAL (T) –MARINE (M)
Particulate inorganic matter River (T to M)
● ●●●●● Lancellotti & Stotz 2004
(mining discharge)
Dissolved metals River (T to M) Vásquez et al. 1999
● ●●●●●
(mining discharge) Vásquez et al. 2000
Particulate inorganic matter River (T to M)
(river floods)
● Miranda 2001
Organic matter Currents Guppy 1906

(carcasses and algae)
●
Arntz & Fahrbach 1991
Organic matter Terrestrial Navarrete & Castilla 1993
vertebrates ● ●●●●● Fariña et al. 2003a
(M to T) Sabat et al. 2003
Nutrients and dead biomass Seabirds Sanchez-Pinero & Polis
(food) (M to T)
● ●●●●●
2000 Ludynia et al. 2005
Between Environments PELAGIC-BENTHIC

POM & phytoplankton Suspension feeders ● ●●● Uribe & Blanco 2001
POM (algal detritus) Currents Bustamante & Branch
● ●●●
1996 Tala & Edding 2005
Between Benthic Habitats NEIGHBOURING COMMUNITIES

POM (detached algae) Currents Rodríguez 2003
● ●●●●●
Tala & Edding 2005
Shell remains Waves and currents Bomkamp et al. 2004
● ●●●●●
Personal observations
Note: POM = particulate organic matter.
& Castro 2003). The transfer of large amounts of algal fragments from subtidal kelp forests toward
the shore has been considered as a principal food source, structuring and maintaining macrofauna
communities on sandy beaches (Colombini et al. 2000, Dugan et al. 2003). Transport of detached
kelp plants or parts to aggregations of sea urchins in tide pools is considered to be an important
trophic subsidy for these grazers (Rodríguez 2003). Arrival of kelp in the intertidal zone of northern-
central Chile continues throughout the year, but highest quantities arrive from late spring until early
autumn, also depending on the proximity to source habitats (Rodríguez 2003). The importance of
kelp transfer to deeper subtidal habitats (for the Californian coast see, e.g., Kim 1992, Harrold
et al. 1998, Vetter & Dayton 1998, 1999) or to the rocky subtidal zone has not been evaluated in
the HCS, but given that the main kelp species are non-buoyant (Lessonia spp.), it is assumed that
large fractions of detached kelp may be accumulating on deeper or wave-sheltered subtidal bottoms.
In addition to kelp detritus, hard-bottom communities also export large quantities of shell
remains to NCs (Bomkamp et al. 2004). Along the coast of northern-central Chile, shell gravel is
relatively common near exposed headlands (Ramorino & Muñiz 1970). These sediments are mainly
composed of shell fragments from barnacles, sea urchins and bivalves, but source habitats, fluxes
of these materials from hard bottoms to sediments and the relevance of local hydrography have
not been examined.
246
Frequency and intensity of exchange processes

Exchange of nutrients or particulate matter among marine communities in northern-central Chile
varies in frequency and intensity (Table 4). Some of the most important and frequent exchange
processes in northern Chile occur in the vertical direction (sedimentation of POM, release of
nutrients into the water column, upwelling of nutrient-rich waters). Horizontal transfer processes
appear to be most intense and frequent in coastal habitats, such as, for example, supply of kelp
detritus to NCs. In contrast to this relatively continuous exchange of material, depositions of
terrestrial sediments to coastal waters or of dead plants and animals to local beaches appear to be
some of the least frequent and unpredictable transfer events. When these events occur, their intensity
is often so high (e.g., Arntz 1986) that they exceed the escape or ingestion capacity of the organisms
in the receiving habitats. This can result in the destruction of local communities and the incorpo-
ration of materials to deeper sediment layers. Transfer of marine-derived materials in colonies of
seabirds and sea lions is also very intense (and frequent), but in northern-central Chile cannot be
utilised by terrestrial organisms due to lack of water. A similar effect is observed in the water
column and sediments of the OMZ where recycling processes are suppressed due to the lack of
oxygen (Graco et al. 2006). Thus the intensity of the fluxes, which overcome the recycling capacity
of receiving communities, may favour the long-term storage of POM not only in shelf sediments
(H.E. González et al. 2004a), but also in terrestrial, intertidal and subtidal habitats of the HCS
(islands with seabird and sea lion colonies, sandy beaches, subtidal kelp accumulations). It appears
to be important to estimate carbon and nutrient export (and storage) not only to shelf sediments
but also to terrestrial soils and intertidal and subtidal bottoms along the HCS.
Propagule supply, dispersal and recruitment variability

Exchange of biological information (i.e., gene flow) depends on the dispersal ability of the organ-
isms in question. Dispersal of individuals determines the scale at which species interact with the
physical environment, the nature and consequences of the interaction with other species, the way
in which they respond to perturbations and ultimately the selective forces and rates to evolve,
speciate or go extinct. Because in most benthic habitats there is a predominance of species with
complex life cycles, which include a free-swimming larval stage (Thorson 1950, Strathmann 1990),
high dispersal capabilities are intuitively associated with most marine organisms. However, this is
not a rule since coexisting with species with planktonic larvae there always exists a myriad of
species with very limited dispersal potential, such as most macroalgae and direct developers or
brooding species (Reed et al. 1992, Kinlan & Gaines 2003, Shanks et al. 2003b). Moreover and to
further complicate things, many species use rafting as an alternative method of long-distance
dispersal (Santelices 1990a, Thiel & Gutow 2005). Despite the early realisation of the high diversity
of life cycles found in every marine habitat, the ecological consequences of such diversity on
species interactions and on the structure and dynamics of benthic communities are only beginning
to be unveiled (Kinlan & Gaines 2003, Leibold et al. 2004, Velázquez et al. 2005).
Methodological approaches to the study of dispersal

The study of dispersal in the ocean is fraught with methodological problems imposed by the
difficulty of following the usually microscopic propagules over extended time. Indirect methods to
estimate aspects of dispersal have been developed. For instance, the use of highly variable neutral
DNA markers offers an unprecedented opportunity to estimate realised dispersal distances (Palumbi
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MARTIN THIEL ET AL.
1995, Kinlan & Gaines 2003, Palumbi 2003, Sotka & Palumbi 2006). Similarly, trace element
microchemistry (elemental fingerprinting) (Swearer et al. 1999, DiBacco & Levin 2000, Zacherl
et al. 2003) or stable isotope ratios (Herzka et al. 2002, Levin 2006) hold promise for identifying
larval origin under specific environmental conditions. However, these techniques do not yet provide
a quantitative measure of the fate of all propagules released from a focal location (i.e., the ‘dispersal
kernel’). Therefore, spatially explicit connectivity among local populations, the type of information
needed regarding the location of MPAs (Botsford et al. 1994, Lockwood et al. 2002, Kaplan 2006),
remains tractable only through the combination of biophysical models linking larval attributes with
advection-diffusion physical processes (Marín & Moreno 2002, Largier 2003, Siegel et al. 2003,
Guizien et al. 2006, Kaplan 2006, Levin 2006, Aiken et al. 2007).
Studies of dispersal in HCS

Studies of dispersal within the HCS are scarce at best. Santelices (1990a) provides a review of
dispersal in marine seaweeds and points out that most information comes from laboratory studies
conducted under idealised hydrographic conditions or from rather anecdotal evidence of colonisa-
tion of new habitats. Studies conducted by incubating seawater samples have demonstrated the
existence of a multispecific ‘spore cloud’ which is present year-round in coastal waters of central
Chile (Hoffmann & Ugarte 1985, Hoffmann 1987). These studies showed the patchy and temporally
variable nature of the spore cloud, but the dispersal distances and mechanisms involved are unclear.
Considering the small size of spores (5–150 µm) and their short duration (a few hours, but it can
be up to few days; Santelices, 1990a), stochastic turbulence diffusion probably plays a major role
in shaping the dispersal kernels in algal dispersal. Nearshore advective currents within the dispersal
scale of spores (e.g., tidal currents, breaking waves, internal tidal bores) cannot be ruled out,
however. Recent studies by Bobadilla & Santelices (2005) conducted by sampling the water column
with a semi-automated sampling device (Bobadilla & Santelices 2004), illustrate the great temporal
variability in multispecific dispersal kernels for major algal groups and dispersal distances exceeding
100 m.
The most direct studies of dispersal of invertebrates in the HCS are restricted to species with
short larval duration, such as tunicates (Castilla et al. 2002a,b, 2004a). These studies sampled larval
distribution at distances from a unique adult population source. Quantitative aspects of dispersal
for species with long-lived larval stages are virtually unknown for any invertebrate or coastal fish
species in the HCS. Several physical processes that can increase offshore and alongshore advection,
or instead increase retention of larvae near shore, have been described for the coast of Chile, such
as upwelling filaments, cyclonic circulation in embayments, topographically controlled eddies and
upwelling shadows and traps (Vargas et al. 1997, Marín et al. 2001, Castilla et al. 2002a, Escribano
et al. 2002, Wieters et al. 2003, Narváez et al. 2004). These features undoubtedly influence dispersal
of coastal species, potentially increasing self-recruitment (Swearer et al. 2002), but their effect on
connectivity among adult populations of any species is hard to demonstrate. A few high-resolution
3-dimensional numerical models of currents in the coastal ocean have been developed and tested
against physical data for different sections of the coast of Chile (Mesías et al. 2001, Aiken et al.
2007). Coupled with Lagrangian larval-tracking techniques these biophysical models can generate
testable hypotheses about dispersal and connectivity in real biological systems (e.g., Aiken et al.
2007).
There is an urgent requirement to develop highly variable, neutral molecular markers such as
microsatellites for algal and invertebrate species inhabiting the HCS system to improve the ability
to infer dispersal distances over ecological timescales and to test hypotheses about connectivity (see
also Population connectivity, p. 252ff.) derived from theoretical models.
248
Settlement studies in the HSC

A more tractable and directly related problem is the settlement and recruitment of species to a
given habitat. Settlement is the process through which a spore or larva makes permanent contact
with the benthic habitat (Keough & Downes 1982). Since most adults of benthic organisms are
sessile or have limited movement, settlement marks the end of the effective dispersal phase. For
brooding organisms with mobile adults or those that use rafting as a secondary dispersal mechanism
this is of course not the case (Thiel & Haye 2006). Recruitment is the input of new individuals to
the benthic population measured at some arbitrary time after settlement. Therefore, while settlement
is expected to reflect the arrival or supply of propagules, recruitment can be substantially modified
by postsettlement mortality. Of the considerable number of studies of supply ecology conducted in
the HCS over the past decades, very few have come close to measuring settlement (Hoffmann &
Ugarte 1985, Hoffmann 1987, Moreno et al. 1993a, 1998, Martínez & Navarrete 2002, Vargas et al.
2004, Lagos et al. 2005, Narváez et al. 2006), and most have actually examined recruitment at
varying time intervals after settlement. The paucity of studies of settlement of marine algae, due
largely to the enormous difficulties of identifying sporelings to species level (Hoffmann 1987,
Santelices 1990a), has not permitted the identification of mechanisms involved in algal settlement
patterns. On the other hand, using intertidal barnacles, mussels and several gastropod species as
model organisms, a few larval transport mechanisms have been demonstrated in the central and
southern HCS. At the locality of Las Cruces in central Chile, which has been characterised as an
upwelling shadow (Kaplan et al. 2003, Wieters et al. 2003, Narváez et al. 2004), the onshore daily
settlement of several invertebrate species is associated with conditions that favour the occurrence
of internal tidal bores, which appear to be common in the area when the water column is well
stratified (Vargas et al. 2004). These results suggest that, for a number of intertidal invertebrates,
internal tidal bores (Pineda 1991, 1994a) can be an important mechanism of onshore transport. In
contrast with results at some sites in the California upwelling ecosystem (e.g., Farrell et al. 1991,
Wing et al. 1995a), studies at Las Cruces showed that settlement of invertebrates was not directly
associated with upwelling-relaxation events, which occur throughout spring and summer over
synoptic timescales. The suggestion here is not that the upwelling-relaxation transport model (e.g.,
Roughgarden et al. 1988, 1991, Wing et al. 1995a,b) plays no role in settlement and recruitment
in central Chile, but rather that at Las Cruces larval transport toward the shore does not seem to
be dominated by these mechanisms. Indeed, spatially intensive studies over a region of about
120 km around Las Cruces showed a clear mesoscale spatial pattern in barnacle settlement, appar-
ently imposed by the topographic variability in upwelling intensity (Lagos et al. 2005). Thus, the
spatial variability in upwelling intensity typical of central Chile (Strub et al. 1998, Broitman et al.
2001, Halpin et al. 2004) might influence the spatial position of the larval pool and probably affects
the scales of dispersal of invertebrates, as suggested also by the study of spatial synchrony in
recruitment of species with contrasting dispersal potential (Lagos et al. in review). Topographic
effects on patterns of settlement and recruitment have also been reported for a variety of brachyuran
crab species (A.T. Palma et al. 2006). Buoyancy fronts produced by river plumes, common from
about 30°S to the south in the HCS, in conjunction with wind stress can also play a role in delivering
larvae to shore (Vargas et al. 2006c). Narváez et al. (2006) also report on the effect of what they
called ‘large warming events’, which occurred a few times in spring–summer in association with
downwelling-favourable (northerly) winds. During these specific large warming events these authors
observed significant synchrony in recruitment of several invertebrate taxa (decapods, gastropods,
polychaetes, mussels and sea urchins), suggesting that larvae could be entrained in these advective
fronts and delivered onshore. A roughly similar phenomenon has been observed around Valdivia
in southern Chile, where southward and onshore movement of warm waters produced by winter
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MARTIN THIEL ET AL.
storms delivered larvae of several gastropod species to the shore (Marín & Moreno 2002, C.A.
Moreno in a personal communication to S.A. Navarrete).
Few studies have directly and simultaneously examined the distribution of larvae in the plankton,
physical processes and settlement onshore in the HCS. Even fewer have examined larval behaviour
under field or laboratory conditions (Poulin et al. 2002a,b, Manríquez et al. 2004, Vargas et al.
2006a).
Patterns of recruitment and benthic communities

Systematic studies of recruitment of species in the HCS have focused on (1) characterising spatial
and temporal variation in the arrival of new individuals for intertidal and a few subtidal species
(e.g., Jara & Moreno 1983, Hoffmann & Santelices 1991, Stotz et al. 1991a, Camus & Lagos 1996,
Martínez & Navarrete 2002); (2) relating these patterns with large-scale oceanographic anomalies,
such as El Niño events (e.g., Moreno et al. 1998, Navarrete et al. 2002); (3) examining the effects
of recruitment variability on population dynamics and recovery of local populations from physical,
biological or human-induced disturbance (e.g., Santelices & Ojeda 1984, Moreno et al. 1993b,
Duarte et al. 1996, Alvarado et al. 2001); (4) determining the consequences of recruitment variation
on the nature and intensity of species interactions in the adult habitat (e.g., Navarrete & Castilla
1990, Moreno 1995, Navarrete et al. 2005, Wieters 2005); and (5) characterising the effects on the
processes that regulate the dynamics of entire intertidal communities over large spatial scales
(Navarrete et al. 2005).
The far-reaching ramifications of persistent variation in recruitment have been most amply
demonstrated in recent studies that quantify patterns of recruitment over large temporal (years) and
spatial (tens to hundreds of kilometres) scales. These studies are starting to shed light on, and find
recurrent patterns in, the causes of the typically large, baffling and usually ‘unpredictable’ variation
in coastal ecosystems. Studies along the California coast have found large-scale regularities in
patterns of recruitment of sessile species that can help reconcile odd experimental results (Menge
et al. 1994, Connolly et al. 2001, Menge et al. 2003). Studies in the HCS conducted by Navarrete
et al. (2002, 2005) have evaluated the effects of variation in wind-driven upwelling on community
regulation along 900 km of coastline between 29°S and 35°S during 72 months. Sharp disconti-
nuities in upwelling regimes around 30–32°S produced abrupt and persistent breaks in the dynamics
of benthic and pelagic communities over hundreds of kilometres (regional scales) (Figure 15A,B).
Rates of mussel and barnacle recruitment changed sharply at 32°S, determining a geographic break
in adult abundance of these competitively dominant species. Analyses of satellite images also
corroborate the existence of regional-scale discontinuities in dynamics and concentration of offshore
surface chl-a that had been previously described at coarser resolution (Thomas 1999, Thomas et al.
2001b). Intertidal field experiments showed that the paradigm of top-down control of intertidal
benthic communities (Castilla & Durán 1985, Paine et al. 1985, Castilla 1999, Navarrete & Castilla
2003) holds only south of this geographic discontinuity. To the north, populations seem recruitment
limited, and predators have negligible effects, despite attaining similarly high abundances. Thus,
geographically discontinuous oceanographic regimes set bounds to the strength of species interac-
tions and define distinct regions for the design and implementation of sustainable management
and conservation policies along the HCS. Further ecological studies using molecular markers are
needed to define the consequences of this variation for the genetic population structure of mussels
and barnacles, as well as for other components of intertidal communities, many of which do not
experience such a discontinuity in recruitment, despite having similar life histories and general
biology (Figure 15C,D).
250
3.5 A Chthamalid barnacles, high zone

3.0
2.5
2.0
1.5
1.0
0.5
0.0
3.0 B Perumytilus purpuratus, mid zone

2.5
2.0
1.5
1.0
Recruits day−1 collector−1
0.5
0.0
2.5 C Semimytilus algosus, mid zone

2.0
1.5
1.0
0.5
0.0
L L N T EL Z TL
M RR UA A L R N N LQ O N S Z
TE A G PT MO MO CU QU TU E TAB IM CIM RU AR P MA P
E C E LC C
70 D Notobalanus flosculus, low zone

60
50
40
30
20
10
ND ND ND
0
L L N T EL Z TL
M RR UA A L R N N LQ O N S Z
TE A G PT MO MO CU QU TU E TAB IM CIM RU AR P MA P
E C E LC C
N S
Figure 15 Average recruitment of intertidal invertebrates along the coast of central Chile at sites ordered
from north to south, from ~29°S to 35°S. Data correspond to long-term (3–7 yr) averages per site of individuals
found in replicated collectors that replaced monthly. The arrow in panels (A) and (B) indicates approximate
position of regional discontinuity in intertidal chthamalid barnacles in the high intertidal zone and the dominant
mussel Perumytilus purpuratus in the mid-zone. See Navarrete et al. (2005) for further details.
251
MARTIN THIEL ET AL.
Population connectivity
Connectivity can be defined as the extent to which populations in different parts of a species’ range
are linked by exchange of larvae, recruits, juveniles or adults (Palumbi 2003) and determines the
degree of cohesion of its genetic pool and the geographic structure of its genetic diversity. The
intensity and geographical scale of connectivity within a species is given by the realised dispersal
through active and passive mechanisms, which depend on species life-history traits and environ-
mental characteristics. Among the numerous dispersal mechanisms reported in marine organisms,
active swimming/crawling and planktonic larval transport, together with rafting and anthropogenic
dispersal, are considered as the most relevant to achieve connectivity among local populations
(Thiel & Haye 2006). Of particular relevance for connectivity of marine species are the temporal
and spatial oceanographic characteristics such as currents, upwelling, water masses and gyres. For
example, even though a species may have long-lived planktonic larvae, in a particular bay the larvae
may not effectively disperse due to local larval retention (e.g., Swearer et al. 1999, Poulin et al.
2002a, Baums et al. 2005). In contrast, benthic species lacking dispersive larval stages can achieve
long-distance dispersal by rafting or anthropogenic transport (Thiel & Haye 2006). Indeed, a recent
study demonstrated that biogeographic patterns along the coast of South Africa are reflected in the
genetic population structure of littoral organisms regardless of their dispersal stages (i.e., with or
without planktonic larvae) (Teske et al. 2006). It seems particularly interesting to pursue this avenue
in the HCS of northern-central Chile where no distinct biogeographic barriers but rather taxon-
dependent breaks exist (see section on biogeography). Moreover, the unique characteristics of the
HCS make it an interesting system to study the genetic connectivity of marine populations. Impor-
tant characteristics of the system for genetic connectivity are its wide geographic extent and the
oceanographic cyclic variations that lead to temporal and spatial changes in population size and
distribution. It is expected that both life-history traits and oceanography play crucial roles in
determining the realised dispersal of marine populations and thus their connectivity and the extent
of their geographic ranges. Few population genetic studies have been published on marine species
of the HCS, although there are several currently being developed on pelagic fishes, marine inver-
tebrates and algae. Nevertheless, some predictions may be formulated and, where possible, validated
through existing examples.
The pattern of genetic connectivity among local populations of a species determines the
geographic structure of its genetic diversity (Figure 16). The frequency, intensity and geographical
scale of dispersal within a species shape the resulting gene flow that counteracts the action of
genetic drift and local selection. In this context, the intensity of genetic drift is principally deter-
mined by population dynamic processes such as population size variation, local extinction, recol-
onisation and founder effects, all intimately related to connectivity among populations. Therefore,
acting both on gene flow and genetic drift, the different patterns of connectivity should result in
different geographic structuring of the genetic diversity. Very high gene flow (at the scale of the
geographic distribution of the species) leads to genetic homogeneity among local populations
independently of the geographic distance. With lower levels of gene flow, different patterns of
population genetic structure may result depending on the association between gene flow and
geographic distance. If the magnitude of the gene flow is associated with geographic distance,
which may be the case for many organisms with planktonic larvae, a genetic cline may form through
the range of distribution, characterised by genetic differentiation being proportional to geographic
distance, a pattern known as isolation by distance (IBD). This pattern will also be influenced by
the direction and strength of the currents. If the magnitude of the gene flow is not strongly associated
with geographic distance, which may be the case for organisms that disperse through passive
mechanisms, the resulting pattern may be chaotic patchiness. So far two parameters have been
252
Gene flow discontinuity

− (barriers to gene flow) +
−
w
e flo Cline Cline + break
gen
f
to
o un Chaotic
Chaotic
patchiness
Am patchiness
+ break
+
+
IBD Homogeneity Homogeneity + break

with geographic
Associated
distance
no
IBD Homogeneity Homogeneity + break
−
Figure 16 Summary of the patterns of genetic connectivity that may result from the interaction among
intensity of the gene flow, its association with geographic distance and its geographic and temporal continuity.
considered: amount of gene flow and association with geographic distance. A third relevant param-
eter is the geographic and temporal continuity of the gene flow, from very continuous to a highly
discontinuous gene flow that will lead to a break in the geographic structure of the genetic diversity.
However, because the amount of time required to reach equilibrium between migration and drift
is at least hundreds of times the generation time of a species, the genetic structuring may also
reflect historic connectivity. Moreover, such equilibrium cannot be reached under high temporal
variability in the pattern of connectivity.
Population connectivity studies in the HCS

A first and very general prediction for the HCS is associated with the long and continuous extent
of the southeastern Pacific coast, without apparent geographic breaks. In this context, IBD and
genetic homogeneity should be the prevalent patterns of geographic structure of the genetic diversity,
particularly for organisms that achieve high gene flow through long-lived planktonic larvae or
frequent rafting routes. The hairy edible crab Cancer setosus, which is of commercial interest, may
represent an example of the above scenario. Gomez-Uchida et al. (2003), using allozymes and
AFLPs (amplified fragment length polymorphisms), show genetic homogeneity over 2500 km of
the Chilean coast for this species. The authors propose that this pattern may reflect the long-lived
larvae (60 days) of C. setosus, the absence of geographic barriers and the oceanographic conditions
(north and southward currents) of the area that allow effective mixing of larvae. A similar pattern
of genetic homogeneity has been observed in pelagic fishes such as Chilean hake (Merluccius gayi
gayi) between 29°S and 41°S (Galleguillos et al. 2000) and Chilean jack mackerel (Trachurus
murphyi) between 20°S and 40°S (E. Poulin unpublished data). It is predicted that species with
high connectivity and extensive geographic ranges may appear less affected by the oceanographic
cyclic variations of the HCS, either because they suffer less population reduction or because they
have a relatively rapid recovery after a disruptive event. Overall, these taxa may lose less genetic
variability and show a faster ecological recovery after ENSO events.
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MARTIN THIEL ET AL.
Conversely, it is predicted that taxa with low dispersal potential will exhibit pronounced genetic
structure and will be the most affected by the oceanographic variations. Genetic studies of the
macroalga Lessonia nigrescens show that gene flow is limited among nearby populations (Martínez
et al. 2003, Faugeron et al. 2005). Additionally, these authors found that 20 yr after the EN
1982–1983 event, which caused a massive mortality of L. nigrescens on 600 km of the coastline,
northward recolonisation had only advanced 60 km (Martínez et al. 2003). Lessonia nigrescens is
a good example of a species very vulnerable to oceanographic changes, specifically EN, and that
may be continuously recovering from drastic population reductions and local extinction, never
reaching migration–drift equilibrium. The genetic structure found for L. nigrescens corresponds to
chaotic patchiness at a small geographical scale (tens of metres), reflecting recent population
dynamic processes (years to tens of years) and life-history traits such as very low distance dispersal
of propagules (Faugeron et al. 2005). Other species that show genetic differentiation at a small
spatial scale are the alga Mazzaella laminarioides (Faugeron et al. 2001) and the edible and
overexploited snail Chorus giganteus that has a low larval dispersal potential (Gajardo et al. 2002).
For the edible and also overexploited scallop Argopecten purpuratus, Moragat et al. (2001) found
both genetic and morphological differentiation between populations at the two protected sides of
the Mejillones Peninsula (50 km apart) and discuss that it is probably due to currents that restrict the
gene flow between the two localities.
It can further be predicted that biogeographic breaks will reflect strong barriers to dispersal
and thus gene flow for species with low dispersal potential, leading to breaks in the geographic
structure of the genetic diversity of species. It has been shown that along the northern Chilean
coasts, habitat discontinuities can cause gene flow interruptions (e.g., Faugeron et al. 2001, 2005).
Species with lower dispersal potential will be more vulnerable to breaks, while species with high
potential of dispersal may not show evidence of a genetic break associated with a biogeographic
break, as is the case of Cancer setosus (Gomez-Uchida et al. 2003). Even though for the HCS it
has not yet been demonstrated that recognised biogeographic breaks correspond with the geographic
distribution of the genetic diversity, it has been shown to be the case for other biogeographic regions
such as Point Conception in the California Current System (e.g., Burton 1998, Wares et al. 2001).
Rafting may be a very advantageous dispersal mechanism for populations that suffer recurrent
extinctions and recolonisations, mostly in the extent of the HCS where macroalgae with high
floatability are very abundant. Once organisms are in a raft that has the potential to be in voyage
for weeks or months, the rafting-mediated gene flow resulting may not be strictly associated with
geographic distance and the resulting pattern of connectivity will depend on the intensity of gene
flow, that is, if the rafting route is frequent, intermittent or episodic (see Thiel & Haye 2006). We
predict that given the abundance of floating macroalgae, rafting routes along the Chilean coast may
be intermittent or frequent, leading to patterns of genetic diversity ranging from chaotic patchiness
to homogeneity. Ongoing studies of the isopod Limnoria chilensis may contribute to the validation
of this prediction. These organisms have the potential to persist in rafts for long periods of time
because they are brooders, have local recruitment and feed on the raft. It is interesting to mention
that even though Lessonia nigrescens shows high genetic differentiation even at small spatial scales,
the geographic distribution of the genetic diversity does not follow an IBD pattern, suggesting that
some long-distance dispersal may occur, although it is not known whether it could be via free-
living spores or on drifting fragments of mature thalli (Faugeron et al. 2005).
The HCS appears to be an interesting model for studying marine connectivity patterns in
variable environments. Despite the general lack of such studies, recent and still unpublished results
on pelagic fishes such as anchovies and sardines, and commercially exploited benthic marine
invertebrates like the gastropod Concholepas concholepas and the bivalve Mesodesma donacium,
support the existence of genetic homogeneity at large geographical scales as a consequence of the
absence of contemporary biogeographical barriers along the HCS for species with high dispersal
254
potential. In general, it is expected that further studies of different biological systems will show
that all the patterns of connectivity (Figure 16) are present along the HCS as a result of the
interaction of present and past environmental conditions with species life-history traits.
Biogeography
Large-scale patterns in the HCS
The pioneer work by S.P. Woodward in 1856, which is probably the earliest biogeographical
classification involving the southeast Pacific (Camus 2001, Harzhauser et al. 2002), was followed
by a series of foundational studies (e.g., Dall 1909, Ekman 1953, Stuardo 1964, Viviani 1979,
Santelices 1980, Brattström & Johanssen 1983, among others) that provided a consistent view of
the major biogeographic features of the HCS temperate area (south of the tropical Panamian
Province), based on physical gradients and patterns of endemism, richness and spatial turnover of
species, and supported by subsequent studies (see reviews by Fernández et al. 2000 and Camus
2001). Overall, two main biotic replacements along the coast differentiate three biogeographical
units (see Brattström & Johanssen 1983 and Camus 2001 for reviews on available classifications):
(1) a warm-temperate biota extending from northern Peru (4–6°S) toward a variable, taxon-
dependent limit in northern Chile (usually 30–36°S), often designated as Peruvian Province, and
dominated by subtropical and temperate species; (2) a cold-temperate biota (also present in southern
Argentina) extending along the fragmented coast of the Chilean archipelago from 54°S to about
41–43°S, corresponding to the Magellanic Province dominated by subantarctic and temperate
species, exhibiting reduced wave exposure and an estuarine condition due to the dilution caused
by high rainfall levels, glaciers and rivers (Ahumada et al. 2000); and (3) a transition zone between
both provinces, characterised by strong numerical reduction of subtropical and subantarctic species
at its southern and northern borders, respectively, rather than by diffusive overlap of biotas. However,
many species occurring throughout this transition zone have a subantarctic affinity and a wide
distribution in Chile (e.g., Menzies 1962, Castillo 1968, Alveal et al. 1973, Santelices 1980),
probably facilitated by the HCS transporting cool water masses toward the north, which is also
considered to be the main reason why the area lacks a definite biogeographic character.
Traditionally, the important physical changes around 42°S are considered to be external forcings
that act as effective filters for dispersal, and with few exceptions, this zone represents the steepest
induced transition along the HCS coast. Contrastingly, the northern limit of the transition zone is
remarkably diffuse for the whole coastal biota (Figure 17) and highly variable depending on the
taxon examined (Camus 2001), which has been attributed so far to the apparent absence of major
physical discontinuities between northern Peru and Chiloé Island (e.g., Brattström & Johanssen
1983, Jaramillo 1987). Such variation mirrors a typical pattern of transitions (Brown & Lomolino
1998), due to differential attenuation rates among taxa related with their different dispersal ability
and physiological tolerance. In fact, some particular taxa (e.g., peracarid crustaceans; Thiel 2002)
show a well-defined overlap of northern and southern species with a gradual replacement pattern.
On a wider taxonomic basis, however, the breaking points for different taxa do exhibit some
latitudinal scattering throughout northern Chile, but they are significantly concentrated around 30°S
and 33°S (see comparative analyses of animal and macroalgal taxa in Brattström & Johanssen
1983, Lancellotti & Vásquez 2000, Meneses & Santelices 2000, Santelices & Meneses 2000, Camus
2001). Notably, these multiphyletic breaks include southern and northern limits of species with
very different life forms and ecological requirements, even involving pelagic groups (e.g., Antezana
1981, Hinojosa et al. 2006). This information strongly suggests that such breaks are not a passive
outcome of dispersal and tracking of key environmental variables. For instance, recent work shows
that latitudinal patterns of SST (the main causal factor invoked in most studies) fail to explain
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MARTIN THIEL ET AL.
PERU
Iquique −20°
Antofagasta
−25°
Southern latitude
−30° Coquimbo −30°
−33°
CHILE
−35°
Concepción
−40°
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30
Figure 17 Break points of distribution along the north-central Chilean coast documented in 29 biogeograph-
ical classifications published between 1951 and 2006 (breaks outside the range 17–38°S are not shown). The
figure includes a wide spectrum of animal and macroalgal taxa of varying hierarchical levels, life forms and
habitats, and some classifications may partially overlap in one or more of these categories. Each classification
is represented along an imaginary vertical axis in correspondence with the numbers at the bottom, where
dashed lines denote the zones in which break points were found (each one marked by a thick horizontal dash),
ranging from one to three break points per study. Horizontal lines highlight the latitudes 30°S and 33°S where
break points tend to concentrate. Classifications (for numbers 1–10 and 11–22, see references in compilations
by Brattström & Johanssen 1983 and Camus 2001, respectively): 1, Mollusca (Carcelles & Williamson 1951);
2, Foraminifera (Boltovskoy 1964); 3, Mollusca (Marincovich 1973); 4, several animal taxa (Knox 1960);
5, eulittoral organisms (Hartmann-Schröder & Hartmann 1962); 6, intertidal Mollusca (Dell 1971); 7, benthic
animals (Semenov 1977); 8, several animal taxa (Viviani 1979); 9, Anthozoa (Sebens & Paine 1979); 10,
several animal taxa (Brattström & Johanssen 1983); 11, benthic macroalgae (Santelices 1980); 12, Bryozoa
(Moyano 1991); 13, fishes (Mann 1954); 14, Isopoda (Menzies 1962); 15, several animal taxa (Ekman 1953);
16, several animal taxa (Balech 1954); 17, planktonic Euphausiids (Antezana 1981); 18, Asteroidea (Madsen
1956); 19, macroalgae (Alveal et al. 1973); 20, sandy beach Isopoda (Jaramillo 1982); 21, Porifera Demo-
spongiae (Desqueyroux & Moyano 1987); 22, several invertebrate taxa (Lancellotti & Vásquez 1999); 23,
demersal fishes (Sielfeld & Vargas 1996); 24, littoral Teleostei (Ojeda et al. 2000); 25, Phaeophyta (Meneses
& Santelices 2000); 26, Rhodophyta (Meneses & Santelices 2000); 27, several animal and algal taxa (Camus
2001); 28, Peracarid crustaceans (Thiel 2002); 29, benthic Polychaeta (Hernández et al. 2005); 30, pelagic
barnacles (Hinojosa et al. 2006).
variations of mollusc diversity along the HCS, which would be determined by shelf area (Valdovinos
et al. 2003), while the distribution of some littoral species appears more related to regional variations
in temperature trends (Rivadeneira & Fernández 2005). Thus, the transitions in northern Chile are
256
not readily explained simply by the contact between warm and cold biotas, and proper explanations
will require a multivariate, integrative approach and an exploration of possible external forcings.
The role of past and present processes in northern-central Chile (18–35°S)

Different lines of historical and ecological evidence suggest that northern-central Chile constitutes
a very complex and dynamical biogeographic scenario. Clearly, present-day patterns are not
divorced from physical changes related to the origin and installation of the cold HCS during the
Tertiary or from subsequent Quaternary fluctuations (e.g., see Villagrán 1995, Hinojosa & Villagrán
1997, Villa-Martínez & Villagrán 1997, Maldonado & Villagrán 2002). The establishment of the
HCS had involved both the northward advance of the subantarctic biota and the northward retraction
of a former tropical/subtropical biota (Brattström & Johanssen 1983, Camus 2001), with conse-
quences that may persist until the present, reflected in the heterogeneous character of the northern
biota. For instance, 10 of the nowadays most common bivalves in northern Chile exhibit upper
thermal tolerances exceeding the highest temperatures recorded during EN events in the past century
(Urban 1994), which is unexpected for species evolving in a cold upwelling system. At least one
of them (Argopecten purpuratus) is thought to be a relict of the Miocene tropical/subtropical fauna
(Wolff 1987). Such physiological ‘anomalies’ suggest the presence of an inertial faunistic component
within the modern northern biota (i.e., remnants of the former warm fauna that escaped the forced
retraction to lower latitudes, and maintained their warm-water characteristics facilitated by recurrent
post-Miocene warming events such as EN) (Wolff 1987). In comparison, the marine flora appears
more homogeneous and dominated mainly by subantarctic species, while tropical/subtropical spe-
cies are virtually absent (Santelices & Meneses 2000). For instance, some common and ecologically
important kelp species are not only more sensitive to warming episodes but also their upper thermal
tolerance varies in accordance with the thermal latitudinal gradient (Martínez 1999).
In this regard, the northern fauna underwent repeated distributional alterations in the past
associated with climatic fluctuations. Some of them involved the simultaneous range retraction and
expansion of different species (e.g., Ortlieb et al. 1994), but more often the occurrence of tropical/
subtropical fauna related to warming (EN-like) events in the Pleistocene (e.g., Ortlieb 1995) and
Holocene (e.g., Llagostera 1979). Moreover, Neogene processes related to the establishment of the
modern upwelling system in the HCS (e.g., shallowing of the OMZ) provoked a mass extinction
of bivalve molluscs (>75% of species), with lasting impacts on their current distribution patterns
and biological characteristics (Rivadeneira 2005), and similar effects have been suggested for the
polychaete fauna (R.A. Moreno et al. 2006a). Notwithstanding, the causal relationships between
historical events and current distribution patterns in northern Chilean waters remain largely unex-
plored, although their importance may be overwhelming.
On the other hand, modern processes also have strong influences in northern Chile, particularly
interannual fluctuations related to ENSO, which, however, should be looked at retrospectively,
acknowledging the frequency and importance of EN-like events throughout the Holocene (e.g, see
Maldonado & Villagrán 2002). Strong EN events can modify the taxonomic composition of littoral
communities (e.g., Arntz 1986, Castilla & Camus 1992, Camus et al. 1994, Vega et al. 2005,
Vásquez et al. 2006) and the geographical occurrence of many species including key structural
components such as intertidal and subtidal kelps (e.g., Lessonia nigrescens, L. trabeculata and
Macrocystis pyrifera). Short-term modifications of community composition during EN occur
through local extinctions and invasions, depending also on local conditions, which either favour or
prevent their occurrence (Arntz 1986, Camus et al. 1994, Martínez et al. 2003, Castilla et al. 2005a).
Moreover, the impacts on key species may scale up to produce long-term biogeographic changes,
as exemplified by the dramatic effects of the 1982–1983 EN event on the intertidal kelp Lessonia
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MARTIN THIEL ET AL.
nigrescens (for similar cases in subtidal kelp beds see Vega et al. 2005, Vásquez et al. 2006 and
the discussion of EN effects here). Lessonia nigrescens plays a key role in Chilean rocky shores
(e.g., Ojeda & Santelices 1984, Castilla 1988, Santelices 1990b), and its presence/absence has
direct effects on community organisation and diversity. The kelp suffered a regionally correlated
local extinction (also involving the loss of its rich holdfast fauna) along 600 km of coastline, which
left a few and highly isolated patches, provoking a strong alteration of its geographical population
structure (Camus 1994b, Martínez et al. 2003). The regional recovery process of L. nigrescens was
slow, more effective toward higher latitudes, and only partial as it failed to re-establish populations
in northernmost Chile (Castilla & Camus 1992). Twenty years later, northward recolonisation
advanced less than 60 km, and some recovered populations lost >50% of their genetic diversity
exhibiting significant IBD (Martínez et al. 2003). These extinctions also lead to transient changes
in biotic interactions within the community (see El Niño section), which had negative effects on
local kelp recruitment and contributed to its slow recovery (Camus 1994a). Additionally, Scurria
scurra, a limpet living on the stipes and holdfasts of Lessonia nigrescens (Muñoz & Santelices
1989), suffered a concomitant extinction. In southernmost localities, Scurria scurra recolonised
and re-established its association with the kelp within 1 yr following the EN event, but in some
northernmost localities it failed to do so for at least 11 yr (Camus 1994b) (and remains absent in
some places until now; P.A. Camus unpublished data). Overall, the ecological, biogeographical and
evolutionary consequences derived from the recurrent extinction-recolonisation dynamics under-
gone by different species in northern Chile are not yet fully understood. However, it may be argued
that they promote changes in spatial patterns of genetic diversity and gene flow, increase between-
community diversity, and affect the dynamics of endpoints of distribution, leading to unstable
biogeographical limits (Camus 2001, Thiel 2002). This last effect can be reinforced by the transient
or permanent invasion of warm-water species favoured by EN episodes (e.g., Soto 1985, Tomicic
1985, Arntz 1986, Castilla et al. 2005a, Coloma et al. 2005, Arntz et al. 2006), thus contributing
to the mixed biogeographic character of the northern Chilean biota. However, while some general
conclusions can be drawn at this level, a proper understanding of large-scale patterns will need to
distinguish their historical and ecological components and consider the physical-biological coupling
generating differential responses among taxa. In this regard, the factors affecting dispersal and
recruitment deserve special attention. Even though EN is known to be related in varied ways to
the recruitment of coastal species (e.g., Soto 1985, Glynn 1988, Vega et al. 2005), in northern Chile
its effects on dominant littoral species may be negligible or highly specific, with no clear association
with interannual variations (Navarrete et al. 2002). Both mesoscale and regional factors related to
the spatial structure of upwelling dynamics seem promising to explain such recruitment variations
(e.g., Lagos et al. 2005, Navarrete et al. 2005). Additionally, the spatiotemporal dynamics of the
OMZ (e.g., Morales et al. 1999, Palma et al. 2005) and the mesoscale eddy activity bounding coastal
ecosystems (Hormazábal et al. 2004) may both play a significant role in understanding the dynamic
connection between oceanographic processes and biogeographic patterns.
El Niño-La Niña in coastal marine communities

The El Niño Southern Oscillation (ENSO) is the largest modern source of interannual variability
in the ocean–atmosphere system (e.g., Wang et al. 1999) and, even though its effects are stronger
in the tropics, it significantly affects marine life in northern Chile. The ENSO cycle has been a
crucial factor in the global climate for at least the past 130,000 yr (Cane 2005), showing continuous,
although variable, activity during the last 12,000 yr (Moy et al. 2002); regionally, it has had a major
influence on the Chilean coast since the Holocene (e.g., see Ortlieb et al. 2000, Maldonado &
Villagrán 2002; see also Biogeography, p. 255ff.). This suggests that coastal communities in
northern Chile have continuously been shaped by impacts of EN events (Camus 1990, 2001).
258
Properties of coastal waters in northern Chile are primarily driven by remote equatorial forcing,
which can provoke strong changes in PP due to the availability of nutrients and essential trace
elements, corresponding to ENSO cycles (Takesue et al. 2004). Such alterations can trigger a
complex chain of biological effects derived from bottom-up controls and physiological constraints,
which may involve several levels of biological organisation at different spatial scales, during and
between EN.
The dramatic and widespread impacts of EN 1982–1983 on coastal communities of northern
Chile allowed the identification of biological changes associated with ENSO such as bathymetric
or latitudinal migrations, invasion by exotic species, behavioural alterations, reproductive and
recruitment failures, increasing population abundance, population decrease due to mass mortality,
and in the most severe cases local population extinctions (e.g., see Soto 1985, Tomicic 1985, Arntz
1986, Glynn 1988, Camus 1990, Castilla & Camus 1992, Sielfeld et al. 2002, Vega et al. 2005,
Arntz et al. 2006, Vásquez et al. 2006). As a whole, these impacts affect all kinds of taxa and
environments, although with clear species- or site-specific components (e.g., in northern Chile none
of these effects involves either an entire taxonomic group or the whole suite of species from a
given place; e.g., see Soto 1985, Tomicic 1985). Moreover, the type and magnitude of impacts, as
well as the range of affected taxa, may vary from one event to another, depending both on the
strength of the event and the type of physical-biological couplings that may take place (e.g., see
Navarrete et al. 2005). On the other hand, some biotic modifications may occur, or have simulta-
neous effects, at both local and regional scales (Camus 1994a, 2001, Vega et al. 2005, Vásquez
et al. 2006), as observed also in the northeast Pacific (Edwards 2004). Additionally, from a socio-
economic perspective, the increase or decrease in abundance or diversity of fisheries resources at
some places may be certainly interpreted as positive or negative effects, respectively (e.g., Arntz
1986). However, from an ecological point of view, it would be as yet uncertain to qualify such
changes in the same terms, even for species with no recognisable importance with variations that
may have unknown or unpredictable consequences for the community.
Thus, simple generalisations on the ecological impacts of EN in northern Chile may still be
inappropriate, except at very specific levels. This situation is mainly because of (1) the lack of
long-term and systematic biological observations encompassing several events, preventing robust
comparisons before, during and after EN conditions, and (2) the irregularity of ENSO itself (e.g.,
Wang et al. 1999) and the lack of correlation between EN and LN in their strength and duration
(e.g., Kerr 1999). Nonetheless, ENSO impacts are indisputably relevant in the ecology of coastal
communities in northern Chile.
One of the key aspects needed to understand EN effects is its recurrent impact on ‘engineer
species’ (sensu Jones et al. 1994) such as kelps, which play a crucial role for the diversity,
complexity, structure and functioning of coastal communities along the southeast Pacific (Graham
2004, Vega et al. 2005, Vásquez et al. 2006). Local extinction of kelps is frequent during strong
EN events (Camus 1994a,b) such as the 1982–1983 episode, when intertidal populations of Lessonia
nigrescens and Macrocystis integrifolia disappeared from the area between 10°S and 21°S and so
did the invertebrate community associated with their holdfasts (Soto 1985; see also Biogeography,
p. 225 ff.). Concurrent and dramatic impacts were reported during the same event (Soto 1985),
affecting ecologically important species of ascidians (e.g., Pyura chilensis), seastars (e.g., Stichaster
striatus, Heliaster helianthus) and several fish species, most of them associated with kelp beds.
However, the implications of these impacts, both at population and community levels, remain
largely unknown.
A long-term series of subtidal community dynamics during variable ENSO conditions
(1996–2005) has been recently published (Vásquez et al. 2006). Although the EN 1997–1998 was
catastrophic and produced local kelp extinctions on the coasts of Chile and Peru (Fernández et al.
1999, Godoy 2000, Martínez et al. 2003), site-dependent conditions allowed the persistence of
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MARTIN THIEL ET AL.
some kelp assemblages of Macrocystis integrifolia and Lessonia trabeculata around 24°S (Martínez
et al. 2003, Vega et al. 2005). These effects would be related to the frequency and intensity of local
coastal upwelling (González et al. 1998, Lagos et al. 2002), which minimised the impact of warming
and retained high concentrations of nutrients within the coastal environment (Takesue et al. 2004).
A long-term analysis of the structure and organisation of kelp communities in northern Chile
(Vásquez & Vega 2004b, Vásquez et al. 2006), which included EN and LN events, showed that the
abundance of Macrocystis integrifolia (1) increased significantly during EN 1997–1998, (2)
decreased during LN 1999–2001, dropping nearly to zero in 2000, and (3) became reestablished
and recovered during a period of positive thermal anomalies in 2002–2003 (Figure 18). This pattern
Figure 18 (A) Upwelling index (Offshore Ekman Transport OET), (B) ENSO index (Southern Oscillation
Index SOI-grey line and Multivariate ENSO Index MEI-black line), (C) temporal variability of Macrocystis
integrifolia, (D) Lessonia trabeculata, (E) benthic grazers, and (F) benthic predator densities, including El Niño
1997–1998 and La Niña 1999–2000.
260
was different from that recorded on the California coast, where the rapid recovery of M. pyrifera
following EN 1997–1998 was favoured by the establishment of a cold period (1998–2000) and the
survival of sporophytes in deep environments (Ladah et al. 1999; Edwards 2004). In northern Chile,
the recolonisation rate of kelp assemblages occurred comparatively slowly (Martínez et al. 2003;
see also Population connectivity, p. 252ff. and Biogeography, p. 255ff.), even though cold conditions
prevailing during 1998–2000 enhanced the upwelling effect. In this regard, the slow recovery of
Lessonia nigrescens after EN 1982–1983 (Castilla & Camus 1992) appeared more related to biotic
constraints: recruitment was strongly reduced by a combination of postsettlement grazing and
inhibition by encrusting coralline algae, while erect coralline algae played a key role as facilitators,
allowing the kelp some escape from grazers and space competitors (Camus 1994a).
On the other hand, the decreased abundance of Macrocystis integrifolia was caused by a
significant reduction in the adult plant population and the lack of recruitment of juvenile sporophytes
(Figure 18). Thus, the disappearance of the M. integrifolia population occurred 2 yr after EN
1997–1998 and was inversely correlated with a population increase of the sea urchin Tetrapygus
niger (Figure 18). In contrast, information from other areas of the southeastern Pacific during EN
1997–1998 showed that superficial warming decreased the abundance of kelp on shallow bottoms,
inducing migrations of grazers to deeper zones (Fernández et al. 1999, Godoy 2000, Lleellish et al.
2001). In northern Chile, during EN 1997–1998 and LN 1998–2000, different events favoured the
increase of sea urchin populations during the cold phase, including (1) induction of mass spawning
due to increases in SST and persistence of upwelling events, (2) reduction in density of adult
seastars, and (3) changes in the feeding behaviour of the seastar Heliaster helianthus, one of the
most important benthic predators on Chilean and Peruvian coasts (Tokeshi & Romero 1995b,
Vásquez et al. 2006) (Figure 18). Thus, the long-term study of subtidal communities suggests that
different bottom-up and top-down factors might control ecosystem changes in northern Chile,
including (1) the intensity and frequency of upwelling, which may buffer the positive thermal
anomalies of SST and maintain high nutrient levels, favouring kelp persistence during EN events;
(2) site-dependent oceanographic conditions, which may generate optimal conditions for spawning,
larval development, and recruitment of echinoderms during and/or after EN events; (3) an overall
abundance increase of carnivores which is correlated with an abundance decline of the most
conspicuous grazers; (4) population dynamics of adult seastars and sea urchins which may become
decoupled during EN events; (5) species-specific population dynamics of some predator species
(e.g., Luidia magellanica), and changes in dietary composition in others (e.g., H. helianthus), which
may promote population increase of its prey, the urchin Tetrapygus niger, during EN events; and
(6) changes in abundance of T. niger, which might be a key factor controlling the development of
two alternate states: environments dominated by kelp beds versus barren ground areas.
In a wider context involving both subtidal and intertidal environments, EN impacts can be
summarised as a large-scale bottom-up effect influencing various (and as yet difficult-to-predict)
levels of marine food webs. However, this is just the initial path for most impacts, and top-down
effects should not be neglected (e.g., see Nielsen & Navarrete 2004). Future research on EN impacts
could consider at least five aspects related to the variability of biological effects, which may serve
as guidelines or study framework: (1) the southward intensity attenuation of EN signals produces
a latitudinal impact gradient, with reduced effects toward higher latitudes (e.g., Castilla & Camus
1992, Martínez et al. 2003); (2) in the spatiotemporal context, many effects are episodic and/or
local (e.g., abundance variability), and some others may propagate their effects to larger spatial
scales (e.g., distribution changes, local extinctions), being highly persistent over time (e.g., see
Camus et al. 1994); (3) on a taxonomic basis, some taxa are recurrently affected (e.g., kelps), others
exhibit no significant impacts (e.g., chlorophytes), and some taxa can be more affected in their
reproduction while others in their recruitment (e.g., Camus 1994a, Navarrete et al. 2005, Vásquez
et al. 2006); (4) the genetic and evolutionary consequences of recurrent phenomena such as mass
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mortalities, extinction-recolonisation processes, and variations in population connectivity are pre-

sumably of critical significance, but they are just beginning to be explored (e.g., see Martínez et al.
2003; see also Population connectivity on p. 252ff. and Biogeography, p. 255ff.); and (5) interannual
variations related to EN and LN may be strongly related to both small-scale processes such as the
Madden-Julian Oscillation (Madden & Julian 1971) and large-scale processes such as the Pacific
Decadal Oscillation (PDO; e.g., Trenberth & Hurrel 1994, Zhang et al. 1997) or the Antarctic
Oscillation (e.g., Gong & Wang 1999), with possible biological implications for benthic community
dynamics that are virtually unknown.
Up to now, the ecological knowledge of EN impacts on the marine communities from northern
Chile continues to be mainly descriptive, focused on a reduced number of species and places.
Nonetheless, prior studies have shed some light on the wide biological scope and geographical
extent of such impacts and the need for comparative, multiscale and long-term approaches to obtain
meaningful results.
Physiological adaptations of marine invertebrates

Physiological variation is the result of genetic, developmental and/or environmental influences
(Spicer & Gaston 1999). Thus, physiological diversity and adaptations are linked to environmental
characteristics and variability. The understanding of how living organisms function (i.e., their
physiology) is aided by comparing the way different animals deal with environmental constraints
(Schmidt-Nielsen 1997).
Major environmental factors affecting the animal’s physiology are temperature, oxygen and
energy (food) availability. The HCS in northern-central Chile is an interesting scenario for running
physiological studies due to changing environmental characteristics, such as (1) the occurrence of
a latitudinal temperature gradient, (2) extended zones with permanent and/or seasonal upwelling
(cold seawater temperature and low oxygen content), (3) some closed bays with relatively high
temperatures (e.g., Antofagasta Bay) compared with the surrounding areas and (4) the occurrence
of thermal anomalies like ENSO. The HCS is characterised also by the occurrence of oxygen-
minimum waters, where physiological adaptations of organisms should be expected, even of species
from shallower (10–50 m) waters, which may occasionally be confronted with low oxygen con-
centrations (when there is upwelling of oxygen-deficient waters). Surprisingly few studies are
available on physiological adaptations to hypoxic conditions of benthic organisms from the HCS.
One of these studies was the characterisation of the pyruvate oxidoreductase enzymes involved in
the biochemical adaptation to low oxygen conditions in nine benthic polychaetes from the HCS
(González & Quiñones 2000). Pyruvate oxidoreductase enzymes permit the metabolism to produce
adenosine triphosphate (ATP) at high rates under environmental or physiological hypoxic conditions
(Livingstone 1983). Interestingly, these enzymes were found to be more numerous and with different
pyruvate consumption rates in the most abundant and worldwide distributed polychaete species
(Paraprionospio pinnata) (González & Quiñones 2000). Another study of biochemical adaptations
to hypoxic conditions in the HCS was done on two key species (in terms of trophodynamics and
abundance) of this system, the euphausiid Euphausia mucronata and the copepod Calanus chilensis
(González & Quiñones 2002). The enzyme lactate dehydrogenase (LDH, a key enzyme of the
anaerobic pathway) from Euphausia mucronata was two orders of magnitude higher than that of
Calanus chilensis. Higher activities of the LDH indicate higher anaerobic capacities, and this may
enable Euphausia mucronata to conduct daily vertical migration through the oxygen-minimum
layer (see also Zooplankton consumers, p. 214ff). In contrast, low LDH activities restrict Calanus
chilensis to oxygenated waters (González & Quiñones 2002). The importance of the interaction
between oxygen and temperature has been explored in recent studies of the brooding behaviour of
decapod crabs (Baeza & Fernández 2002, Fernández et al. 2006b).
262
Closed bays with relatively high temperatures and productivity in the HCS offer suitable
conditions for the permanence and reproduction of the scallop Argopecten purpuratus, a species
more characteristic of warm waters. An increment of 2.5°C in bottom temperatures (normally
15.5°C) during EN 1982–1983 in Tongoy Bay (30°S) augmented dramatically gonad mass and
spawning, and as a consequence spat (juvenile) collection exceeded levels from previous years by
300% (Illanes et al. 1985). However, total gonadal levels of lipids and proteins increased markedly
in A. purpuratus conditioned for reproduction at 16°C, but these increases were less pronounced
at 20°C (Martínez et al. 2000). Moreover, during gonad maturation muscle carbohydrate levels
dropped considerably, as well as the activity of a pyruvate oxidoreductase, the enzyme octopine
dehydrogenase (Martínez et al. 2000). Muscle carbohydrate (i.e., glycogen) and glycolytic enzymes
have been shown to decrease greatly in other scallop species such as Chlamys islandica and Euvola
ziczac (Brokordt et al. 2000a,b). This leads to a decrease in muscle metabolic capacity and thus in
escape capacities, which is facilitated by muscle contractions. A reduction of escape capacities
during reproduction has been observed in Argopecten purpuratus as well as in Chlamys islandica
and Euvola ziczac (Brokordt et al. 2000a,b, 2006).
In the intertidal and shallow subtidal zones of the HCS, temperature is the main variable
changing over various spatial and temporal scales, with unpredictable interannual patterns. Under
normal conditions, physical environmental conditions are relatively stable in the shallow subtidal
between 18°S and 35°S (HCS), where salinity typically ranges between 34 and 35 and temperature
may vary from 12°C to 22°C. However, due to terrestrial influence, the temperature conditions in
the intertidal are different along this latitudinal gradient. For example, the range of mean temper-
atures registered in high intertidal pools during the summer is ~13–33°C in Antofagasta (23°S),
~13–30°C in Carrizal Bajo (28°S), and ~11–25°C in Las Cruces (33°S) (Pulgar et al. 2006). During
EN, these differences in thermal conditions may be enhanced.
To evaluate phenotypic plasticity or evolutionary responses of organisms to different habitat
temperatures, comparative studies have typically focused on species distributed along latitudinal
gradients (Vernberg 1962, Graves & Somero 1982, Stillman & Somero 2000, Pulgar et al. 2006).
However, local thermal gradients (TGRs) can be formed by fine-scale variation in, for example,
the marine intertidal vertical zones. The intertidal zone is characterised by important spatial and
temporal gradients of temperatures, which may be equivalent to those found over a large latitudinal
gradient. Intertidal organisms have evolved physiological tolerance adaptations that are important
in determining the upper vertical distribution of the species. Studies of congeners or conspecifics
allow adaptive variation to be clearly demarcated, independent of effects of phylogeny (Stillman
& Somero 2000). Crabs of the genus Petrolisthes (Anomura: Porcellanidae) are widely distributed
not only along the intertidal zone of the HCS, but also worldwide, covering huge latitudinal
gradients. One of the few studies of physiological adaptations of marine invertebrates in the HSC
(Las Cruces, 33°S) was done in five species of the genus Petrolisthes (P. granulosus, P. laevigatus,
P. violaceus, P. tuberculatus and P. tuberculosus) (Stillman & Somero 2000). Each species is found
at different vertical levels, from the low (P. tuberculosus), mid-low (P. tuberculatus), middle
(P. violaceus), mid-high (P. laevigatus) to the high (P. granulosus) intertidal. The limits of thermal
tolerance (LT50) were strongly correlated with the vertical position of the species in the intertidal
zone (y = 36.02 − 1.88x, r 2 = 0.97) and with the maximal habitat temperature (Table 5) (Stillman &
Somero 2000). Thus, species have adapted their upper thermal tolerance limits to coincide with
microhabitat conditions. Interestingly, mid-high and high intertidal species (P. laevigatus and
P. granulosus, respectively) live near their limits of thermal tolerance. While these LT50 values offer
some hints, it may be extremely interesting to explore at which temperatures these organisms enter
suboptimal ranges, that is, where they may be able to survive but where growth and reproduction
may be compromised. Petrolisthes laevigatus from southern-central Chile dramatically reduces
oxygen consumption between 18°C and 20°C (maximal average temperature range found in its
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MARTIN THIEL ET AL.
Table 5 Thermal tolerance limits (LT50), and spring maximal habitat temperature of
Petrolisthes along intertidal vertical gradient in a locality at the HCS (33°S)
Vertical position ~ Limit of thermal tolerance ~ Maximal habitat temperature
Species of Petrolisthes in the intertidal (LT50, °C) (°C)
P. tuberculosus Low 27.5 14
P. tuberculatus Mid-low 28.5 18
P. violaceus Middle 30.5 18
P. laevigatus Mid-high 31.5 28
P. granulosus High 35.0 33
Note: Data from Stillman & Somero (2000).
habitat), which suggests the beginning of the organism’s decompensation, or the commencement
of another homeostatic mechanism independent of oxygen consumption (Yaikin et al. 2002). Higher
thermal stress or having thermal limits close to actual maximal habitat temperatures might increase
the ‘cost of living’ of the species in the upper intertidal (Somero 2002, Stillman 2002). This higher
cost of living would be associated with the cost of repairing thermal damage (heat-shock proteins,
Hsp) and adapting systems through acclimatisation (Somero 2002). Although tropical Petrolisthes
species have higher thermal limits than HCS species, the latter show a wider range of thermal
tolerance between low and high intertidal species (Stillman & Somero 2000). Moreover, low intertidal
HCS species show greater phenotypic plasticity in their thermal tolerance than high intertidal species
(Stillman & Somero 2000). Considering global warming, species inhabiting the upper intertidal
zone, living at the ‘edge’ of their thermal limits, would be more affected than species from the
lower intertidal, which live far from their thermal limit and with a greater thermal phenotypic
plasticity. These physiological traits may have an important effect on the borders in the latitudinal
distribution of a species and consequently also on biogeographic limits.
Thermal effects that occur outside the normal physiological range involve deleterious changes
at the cellular level, especially in systems involved with oxygen uptake, delivery and utilisation
(Stillman 2002), such as the cardiac system (Frederich & Pörtner 2000). The heart of a crab species
living in the upper intertidal has an Arrhenius break temperature (ABT, temperature at which a break
occurs in the slope in an Arrhenius plot, i.e., log rate vs. reciprocal of absolute temperature, K)
that is 5°C higher than in a crab species from the low intertidal (Stillman & Somero 1996). These
differences were associated with the Na+ K+ ATPase (adenosine triphosphatase) activity, which is
necessary for the establishment of the membrane action potential that permits the heartbeat (Stillman
2002). Similar to the rate of heartbeat, oxygen consumption by isolated mitochondria exhibits a
‘break’ at some high temperatures (Dahlhoff & Somero 1993). Phenotypic plasticity in the ABT
of mitochondrial respiration has been observed in abalone Haliotis congeners from different thermal
habitats (latitude and vertical positions along subtidal-to-intertidal gradient) (Dahlhoff & Somero
1993). Protein synthesis and heat-shock response has been shown to change spatially among
gastropod (Tegula) congeners from the temperate subtidal to the low intertidal zones (Tomanek
2001) and seasonally in the bivalve Mytilus trossulus (Hofmann & Somero 1995). Hsp have the
function of refolding and ‘rescuing’ proteins damaged by thermal denaturation (Becker & Craig
1994, Hofmann & Somero 1995). Intertidal species of Tegula showed greater expression of Hsp70
than the subtidal species when temperature increases (Tomanek 2001). The energy cost associated
with replacing damaged proteins and maintaining Hsp may be an important proportion of cellular
energy demands (Hofmann & Somero 1995, Somero 2002).
Intertidal and shallow subtidal invertebrates present a ‘cascade’ of physiological responses that
enable them to adapt and finally survive changes in environmental conditions. Despite the important
264
latitudinal and vertical environmental conditions gradient of the HCS, there are few studies of
physiological responses of invertebrates living in this ecosystem. The range of temperatures regis-
tered in the intertidal (rocky pools) along this latitudinal gradient (Pulgar et al. 2006) is very similar
to the TGR from the low-to-high intertidal zone observed in central Chile (Table 5). Therefore, it
could be expected to find similar thermal physiological variability and local adaptations of conge-
ners or conspecifics in the latitudinal gradient to those found in the intertidal vertical gradient.
Because in this area the pattern of environmental variability shifts from a relatively predictable
seasonal pattern to a more unpredictable pattern of high interannual variability (i.e., ENSO),
physiological characterisation of congeners or conspecific organisms inhabiting the intertidal and
shallow subtidal zones along the latitudinal gradient of the HCS would be particularly interesting.
Moreover, since algal availability increases from northern to southern Chile (Santelices & Marquet
1998), and physiological compensation associated with environmental stress increases cost of living
(Somero 2002, Stillman 2002), latitudinal changes in food availability should also be considered
in future studies. It appears particularly interesting to examine how increased costs of living near
the distribution limit of a species influences its reproductive potential.
Reproductive patterns of selected marine invertebrates

in the HCS
Some of the factors that vary with upwelling intensity and persistence, such as temperature and
PP, are known to critically affect per capita reproductive investment of marine invertebrates (e.g.,
MacDonald & Thompson 1985, 1988, Clarke 1987, Brey 1995, Phillips 2002). In the northeastern
Pacific, reproductive hot spots coincide with regions exhibiting high PP, which suggests not only
that bottom-up processes play a central role in explaining reproductive output, but also that spatial
heterogeneity in reproduction needs to be considered in conservation and management plans (Leslie
et al. 2005). The clear break in eddy kinetic energy and equatorward wind stress reported at 30°S
in the southeastern Pacific (Hormazábal et al. 2004) coincides with two contrasting regimes in chl-a
concentration both in coastal areas and offshore (Yuras et al. 2005). Chl-a concentration is nega-
tively correlated with seawater temperature along the HCS (Strub et al. 1991, Thomas et al. 2001b).
The effects of small- and large-scale variation in environmental conditions related to upwelling
persistence and strength on reproductive patterns along the HCS have recently been analysed.
Primary productivity and gonad production

Gonad and egg production of marine invertebrates do not exhibit a clear latitudinal cline in investment
in reproduction along the HCS in central Chile (Fernández et al. 2007). However, gonad production
shows variable patterns throughout the study region, which extends from 28°S to 36°S, and this
variability appears related to the trophic level of the species being investigated and the proximity of
the study sites to upwelling centres. Carnivores such as Concholepas concholepas and Acanthina
monodon do not show variation in gonad investment between 28°S and 36°S (Figure 19). In contrast,
some of the dominant intertidal suspension-feeding species, Perumytilus purpuratus and Nothoch-
thamalus scabrosus, and one of the most abundant herbivores, Chiton granosus, exhibit strong
variation in gonad production among sites, even between adjacent locations (Fernández et al. 2007).
Investment in reproduction of suspension-feeding and herbivore species is constantly higher in
some sites (e.g., Los Molles 32°24′S, Consistorial 33°49′S) and lower in others (e.g., Montemar
32°96′S, Matanzas 33°96′S). Furthermore, there is a positive and significant correlation between
investment in gonads among Perumytilus purpuratus, Nothochthamalus scabrosus and Chiton
granosus (p always < 0.05; P. purpuratus-N. scabrosus: R = 0.89, P. purpuratus-Ch. granosus:
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MARTIN THIEL ET AL.
28°S 30°S 32°S 34°S 36°S
0.15 A
0.1
0.05
0
0.03
B
0.02
Reproductive output
0.01
0
0.125
C
0.02
0.075
0.05
0.025
0
0.5
D
0.4
0.3
0.2
0.1
0
500
E
400
n/basal area
Brood size
300
200
100
0
Huasco
Pta. Choros
Temblador
Cta. Hornos
Arrayan
Totoralillo
Guanaqueros
Los Molles
Montemar
El Quisco
Matanza
Pichilemu
Consistorial
Quintay
Figure 19 Reproductive output of (A) Concholepas concholepas, (B) Acanthina monodon, (C) Chiton granosus,
(D) Perumytilus purpuratus, and (E) brood size (number of embryos per unit of basal area) of Nothochthamalus
scabrosus, along the study site which ranges from 28°S to 36°S in the eastern South Pacific Ocean. The white
bars are used for carnivore species, the black bars for herbivore or filter-feeding species. The white arrows
indicate upwelling centers, the black arrows point to sites not influenced by upwelling along the HCS.
R = 0.64, N. scabrosus-Ch. granosus: R = 0.73; Fernández et al. 2007). Investment in gonads of

carnivore species does not show any correlation with the reproductive patterns exhibited by herbi-
vores or suspension-feeding species. The patterns reported were consistent over the 3-yr study
(coefficient of variation = 1.3%). These results suggest that the small-scale variation in environ-
mental conditions along the upwelling-favourable region of the HCS seems to affect gonad pro-
duction and consequently larval production, and therefore that some locations potentially serve as
source populations of propagules. Transplant experiments support the hypothesis that local environ-
mental conditions determine reproductive output. Suspension feeders (e.g., Perumytilus purpuratus)
266
and herbivores (e.g., Chiton granosus) transplanted between the most and least suitable sites for
reproduction in central Chile showed that reproductive output was strongly determined by the site
to which the animals were transplanted while the site of origin showed a negligible effect (Fernández
et al. 2007). Organisms transplanted to Los Molles showed high reproductive output and organisms
transplanted to Matanzas showed low reproductive output, regardless of the site of origin (Fernández
et al. 2007). These contrasting results suggest that environmental variables, such as PP, may affect
investment in gonads of lower trophic level benthic invertebrates. These environmental conditions
seem to be related to the spatial variation of upwelling conditions. The central coast of Chile is
dominated by seasonal wind-driven upwelling that forces cold, nutrient-rich water into the upper
water column (Wieters et al. 2003). However, it is remarkable that the well-documented relationship
between cold upwelled water and high chl-a concentration over large spatial scales off the coasts
of Chile and California (Strub et al. 1991, Thomas et al. 2001a) is not observed at smaller spatial
scales (Wieters et al. 2003). Between 28°S and 36°S, the lowest chl-a concentrations are associated
with coldest upwelled waters (Wieters et al. 2003). Matanzas and Montemar are sites influenced
by upwelling centres, in contrast with the lower influence of upwelling in areas such as Los Molles
or Consistorial (Broitman et al. 2001, Wieters et al. 2003). Upwelling centres also show higher
growth rate of corticated algae, which are not consumed by herbivores, and low growth of ephemeral
algae (Nielsen & Navarrete 2004). Although temperature decreased from 28°S to 36°S and is lower
in upwelling centres, gonad production is not correlated with seawater temperature (Fernández
et al. 2006a). Most likely, the low chl-a concentration associated with upwelling conditions (Wieters
et al. 2003) and low abundance of benthic ephemeral algae (Nielsen & Navarrete 2004) are the
main factors affecting reproductive output of lower trophic level, intertidal species along the HCS.
Evidence from other geographic regions supports the hypothesis that patterns of PP associated with
upwelling conditions determine reproductive output (Leslie et al. 2005). However, more information
is necessary to clearly identify the set of environmental variables affecting reproductive investment.
Temperature and brooding requirements

Although the effect of temperature on gonad production is not evident along the upwelling region
associated with the HCS between 28°S and 36°S, studies conducted over larger spatial scales (and
wider temperature ranges) suggest that temperature can affect egg production in species that
aggregate embryos. Oxygen is a limiting factor in embryo aggregations of marine invertebrates
(Cohen & Strathmann 1996, Lee & Strathmann 1998, Fernández et al. 2003) and temperature
affects oxygen availability in different types of embryo packing (Brante et al. 2003, Fernández
et al. 2006a). Studies of the brachyuran crab Cancer setosus show that brooding females respond
to the increased oxygen demand of the embryos at higher temperatures by increasing abdominal
flapping frequency, a behaviour that supplies oxygen to the brood (Brante et al. 2003). Similar
patterns of female response to embryo oxygen demand have been reported throughout embryo
development in other crab species (Baeza & Fernández 2002, M. Fernández et al. 2002). The change
in female brooding behaviour (abdominal flapping frequency) produces a higher rate of oxygen
supply, dramatically affecting the costs of brooding. Between 10°C and 18°C, a 45% increase in
brooding costs was estimated for large-size crabs, such as C. setosus (Brante et al. 2003). This
substantial increase in brooding behaviour and cost seems to affect egg production and survival
(Fernández et al. 2003). Field data showed that gonad investment (or reproductive output) in
C. setosus is lower in northern Chile (approximately 20°S) than in central (30–33°S) and southern
(40°S) Chile. Lardies & Castilla (2001) reported a similar latitudinal trend in reproductive output
for Pinnaxodes chilensis. Moreover, embryo loss in Cancer setosus increases with temperature
(Brante et al. 2003), suggesting that temperature also affects embryo survival throughout the
brooding period. Reproductive output of most brachyuran crab species does not vary south of 30°S
267
MARTIN THIEL ET AL.
(Brante et al. 2004). The contrasting pattern in reproduction between populations of brachyuran
crabs north and south of 30°S seems also to occur in other species that aggregate embryos. It is
interesting that despite the lack of difference in reproductive output of Concholepas concholepas
between 28°S and 36°S (Figure 19), embryo packing shows a clear break that coincides with the
break exhibited by brachyuran crabs (north and south of 29–30°S). The mean number of embryos
per unit of capsule area was significantly lower in capsules from sites north of 29°S than south
thereof (20.6 vs. 31.8, respectively), over a range of study sites located between 30°S and 42°S.
Although there is no dramatic break in temperature at 30°S (Broitman et al. 2001), the pattern of
embryo packing is explained by the mean temperature at the study site shortly before egg deposition.
The evidence accumulated to date suggests that the higher brooding costs north of 30°S seem
to affect reproductive output and that the effect is consistent across the brooding modes and taxa
studied so far. This finding suggests that environmental conditions are less favourable for brooding
species in the northern part of the HCS. In fact, large-scale studies of the patterns of species
distribution in relation to larval developmental mode support the hypothesis that the spatial distri-
bution of brooding species is explained by the cost of brooding, which is associated with the cost
of oxygen provision (Astorga et al. 2003, Marquet et al. 2004). It is less clear if the impact of the
cost of brooding on reproductive output may change with adult body size. Recent studies have
shown that small crab species perform the same active brooding behaviours as large species (e.g.,
Pisoides edwardsi, Acanthocyclus gayi; Fernández et al. 2006b). However, mean oxygen consump-
tion of brooding females, which is a proxy of brooding cost, is not significantly different from
oxygen consumption of non-brooding females (Fernández et al. 2006b). These results contrast with
previous studies of large species, showing a 2- to 3-fold increase in oxygen consumption by females
carrying later-stage embryos over non-brooding females (e.g., Cancer setosus: Baeza & Fernández
2002; Homalaspis plana: Ruiz-Tagle et al. 2002; Ovalipes trimaculatus: Fernández & Brante 2003).
Therefore, the patterns described for brooding species may be dependent on adult body size. In
fact, the small crab species Acanthocyclus gayi and A. hassleri do not show any pattern in repro-
ductive output along the region extending from 28°S to 36°S (Espinoza 2006). Although existing
evidence on reproductive output along the HCS suggests that the behaviour of populations north
and south of 30°S depend on the cost of oxygen provision in larger-size species, more studies are
needed in order to generalise the findings on the effects of temperature on brooding costs, the link
between adult size and brooding mode, and the consequences on species distribution, especially in
regions influenced by upwelling where low oxygen conditions cover extended regions of the ocean
(Fuenzalida et al. accepted).
Larval life in the HCS

Oceanographic conditions in the HCS ecosystem expose planktonic larval forms to environmental
conditions in which they do not behave simply as a passive particle. Therefore, morphological,
physiological and behavioural characteristics present in fish and benthic invertebrate larval forms
inhabiting any particular habitat can be interpreted as effective local adaptations evolved to face
this unique ecosystem. Herein, the aim is to give a brief overview of some behavioural and feeding
traits together with transport processes described in larval forms of benthic invertebrates and fishes
inhabiting the HCS.
Behavioural traits
Larvae of many marine benthic invertebrates, differing from lecitotrophic short-lived larvae, spend
extended periods in the plankton prior to settlement and metamorphosis (Thorson 1950, Pechenik
1986, 1999). Usually, during their pelagic phase, planktotrophic larvae (PL) of marine benthic
268
invertebrates must remain suspended, locate and gather food, avoid predators and unfavourable
conditions, disperse to new areas and select sites for settlement (Strathmann 1974, Palmer &
Strathmann 1981, Scheltema 1986). The importance of larval behaviour has been recognised as an
important subject in marine ecology and the recent focus on the role of nearshore oceanographic
processes controlling larval dispersal and recruitment of benthic organisms has revived interest in
the behaviour of larvae in the plankton (reviewed in Le Fèvre & Bourget 1992). Although larval
transport seems to be mainly controlled by hydrographic factors (Thorson 1950), larval behaviour
may influence their final destination (e.g., Butman 1987, Pineda 1994a,b). In general, sources of
mortality in the dispersal phase for PL are food limitation, extreme conditions of temperature and
salinity, low dissolved oxygen, UV radiation and pollution (Pechenik 1986). However, because of
the difficulties associated with larval tracking in the field, the relative importance of each potential
source of larval mortality in nature is still largely unknown. Field and laboratory experiments have
revealed a number of behavioural mechanisms that allow larvae to contend with these selective
pressures on mortality. However, specific studies of characteristics displayed by PL throughout the
HCS are largely absent. Few published studies regarding larval characteristics linked with ecological
importance have been conducted in Chile (e.g., Manríquez & Castilla 2007). Field evidence of a
DVM pattern in competent larvae has been described for the muricid gastropod Concholepas
concholepas (Poulin et al. 2002a,b). It has been suggested that the behaviour of competent larvae
of this species may help them to avoid offshore transport during upwelling events. Studies with
several crustacean species have shown clear synchronisation between reproduction and the hydro-
dynamic processes promoting larval transport or retention (Yannicelli et al. 2006a). Similar studies
with other PL in the HCS incorporating behavioural or physiological responses to environmental
variables such as currents, temperature and salinity are urgently needed. However, along the HCS
those studies are precluded by the absence of basic knowledge such as larval identification in many
species of Chilean invertebrates. Work to investigate the scale of the source–sink dynamics of PL
in the temporally heterogeneous HCS ecosystem is also largely absent. More recent techniques
involving chemical tags in calcified structures of PL have been successfully used to identify potential
larval source and sink areas, and larval trajectories in the HCS (Zacherl et al. 2003).
Feeding and larval food environment in the HCS

Recruitment is recognised as the leading determinant of population dynamics of benthic invertebrate
and fish species and it strongly adjusts the importance and intensity of species interactions in the
HCS (see above). Since PL rely on food to grow and develop, availability and quality of food
particles during larval development are important factors influencing larval recruitment (Vargas
et al. 2006a). Despite the high variability in larval food composition, there are well-documented,
persistent, temporal and spatial differences in plankton structure and abundance along the HCS.
For instance, large and geographically persistent heterogeneity in chl-a has been documented along
the Chilean coast (Thomas et al. 2001b, Yuras et al. 2005; and see Dominant primary producers
and their role in the pelagic food web, p. 210ff.). The general pattern indicates that a large percentage
of chl-a is found in the large phytoplankton fraction in permanent coastal upwelling areas off
northern Chile (Iriarte & González 2004; see also Dominant primary producers and their role in
the pelagic food web, p. 210ff.), as well as during spring/summer at seasonal upwelling sites in
central Chile (González et al. 1989, Vargas et al. 2006b). Similarly, a large and geographically
persistent heterogeneity in chl-a levels has been observed to the north of latitude 32°S (Yuras et al.
2005).
The scarcity of available published data on larval fish and invertebrate feeding makes it difficult
to evaluate potential larval survival and recruitment along the HCS. Currently, the largest body of
literature is available for larval fish of commercially exploited species. Because of their seasonal
269
MARTIN THIEL ET AL.
Autotrophs Heterotrophs −
Picoautotrophs Bacteria
High
Food size
chlorophyll Autotrophic
nanoflagellates Heterotrophic
nanoflagellates
Spatial/temporal heterogeneity
+
Diatoms Microprotozoa
Copepod
stages
mid-size
Physical processes
prey
Affecting both food and larvae
large prey Turbulence
Internal waves, tides
River plume motion
small prey
Frontal structures
Filaments
Upwelling/downwelling
Microprotozoa +
Diatoms
Food size
Autotrophic Heterotrophic
nanoflagellates nanoflagellates
Picoautotrophs Bacteria −
Low Heterotrophs
chlorophyll
Autotrophs
Figure 20 Conceptual scheme of main pathways of interaction in coastal food webs involving invertebrate
(i.e., veliger competent larvae and barnacle nauplii) and fish larvae under spatial/temporal variation in chlo-
rophyll levels in the Humboldt Current System. The thickness of the arrows represents main predator-prey
interactions. The sizes of the boxes or circles represent the dominance in terms of biomass of a specific food
item (both autotrophic and heterotrophic prey) during each condition. Physical processes discussed in this
chapter, which affect both larvae and food distribution, are also included. Arrows directed to food items at
top or bottom boxes were included for convenience.
nature, in the coastal upwelling area off central Chile (36–37°S), the changes in the phytoplankton
protozoan and microplankton community that constitute the food supply for larval fish also occur
on a seasonal basis (Vargas et al. 2006b). Accordingly, larvae produced in the middle of winter in
the HCS, when PP and seawater temperatures are low and wind-induced turbulence in the upper
part of the water column is high, are probably not going to face the same prey as those produced
in summer, when upwelling and PP are at maximum (Figure 20). Most of the studies of larval fish
feeding have focused on changes in prey field, diet overlap, and their influence on larval feeding
over short timescales, between adjacent areas, or have attempted to investigate whether evidence
of starvation occurred in wild-collected larvae (Llanos et al. 1996, Balbontín et al. 1997, Pizarro
et al. 1998, Llanos-Rivera et al. 2004). For the anchovy, Engraulis ringens, in northern Chile
(20–21°S), scarce incidence of starvation was even observed during autumn, a season of reduced
270
plankton production (Pizarro et al. 1998). In northern Chile, the diet of the myctophids Diogenichthys
laternatus and Triphoturus oculeus, which feed diurnally and share the upper water column (0–200
m depth), was shown to vary according to prey availability in the field. The diets of both species
overlap in periods and areas where food is more abundant (e.g., copepods, copepodids, nauplii,
invertebrate eggs and ostracods), but differ under conditions of low prey availability (Rodríguez-
Graña et al. 2005). Other studies in central Chile have shown that either microplankton concentra-
tions did not appear limiting in winter (Castro et al. 2000, Hernández & Castro 2000, Castro 2001)
or larval diet overlap occurred among several species but during periods of high food abundance
(Llanos et al. 1996, Balbontín et al. 1997, Llanos-Rivera et al. 2004). Hence, the few studies carried
out on feeding of larval fish along the HCS in Chile suggest that, although the larval food abundance
may vary among seasons and localities, starvation due to limiting food availability alone does not
seem to be such a common feature, even in seasons of lower production (autumn and winter). For
starvation to occur, other factors may be necessary, such as increased turbulence, which may play
a concomitant role during the low productivity seasons, at least in southern Chile (Cury & Roy
1989, Castro et al. 2002). Scarce dietary overlap and local morphological and physiological adap-
tations (i.e., increased reserves in fish eggs and larger yolk size in fish larvae; Llanos-Rivera &
Castro 2004, 2006) seem to exist in larval periods when ontogenetically food is highly necessary
(i.e., onset of feeding) or when food becomes less abundant in the environment (Balbontín et al.
1997, Llanos-Rivera et al. 2004, Rodríguez-Graña et al. 2005).
For benthic invertebrates, sharp spatial transition in phytoplankton biomass associated with
upwelling dynamics has been assumed to have important effects on larval condition (Wieters et al.
2003). This assumption is based on the idea that phytoplankton is the most important food item
for PL. A major impediment to understanding how food quantity and quality influence larval life
under natural conditions is that virtually all published information comes from laboratory rearing
studies in which larvae are offered a monospecific or controlled-mix algal culture. One of the first
studies done in the HCS analysing feeding preferences in larval invertebrates, by Vargas et al.
(2006a), found evidence that barnacle nauplii (Jehlius cirratus and Notobalanus flosculus) and
veligers (Concholepas concholepas) exhibited high consumption of heterotrophs (i.e., ciliates and
dinoflagellates), but the size spectrum of food particles removed by barnacle nauplii was in contrast
with those for C. concholepas veligers. Barnacle nauplii preyed heavily on small picophytoplankton
(<2 µm), whereas competent C. concholepas veliger larvae were mechanically unable to feed on
small cells, and the highest removal rates were of nano- and microphytoplankton (>20 µm). The
ability of barnacle nauplii to feed on small prey hinges on the small spaces between their limb
setules (Stone 1989). This important finding indicates that omnivorous larval feeding should be the
norm in the pelagic ecosystem and might explain why larvae maintain positive growth in environ-
ments where phytoplankton is thought to be limiting (Crisp et al. 1985). Therefore, the scarce
published information for the HCS suggests that the inference of patterns of larval condition and
recruitment over large scales from chl-a biomass, now easily measured from satellite images (e.g.,
Thomas et al. 2001b), has to be regarded with caution. The scenario suggests that a large spectrum
of food particles is available for larval feeding, and species may adapt their feeding preferences in
relation to temporal/spatial food distribution along the HCS as well as their physiology and energetic
reserves to counteract the spatial and temporal variations in food quality and quantity (Vargas et al.
2006b) (Figure 20).
Upwelling and larval transport processes

The wind-driven seasonal upwelling, besides its paramount effect on PP in the coastal zone, induces
profound changes in the dynamics of coastal waters that directly affect the distribution and abun-
dance of organisms in the nearshore areas as well as over the continental shelf and slope. To reside
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MARTIN THIEL ET AL.
in these hydrodynamically variable but highly productive environments, coastal and offshore
organisms have developed a series of reproductive strategies that enable them to counteract the
effects of offshore advection in the surface Ekman layer or take advantage of other oceanographic
processes to return to the coastal zone (see also Coastal oceanography, p. 205ff.). For instance, on a
seasonal timescale, small pelagic clupeiform fish tend to synchronise their reproductive timing with
processes that induce shoreward transport and coastal retention. Probably the best-documented species
in the HCS of central Chile are the anchovy (Engraulis ringens) and common sardine (Strangomera
bentincki) that reproduce during winter, when the wind-driven Ekman layer is directed shoreward and
thus eggs and larvae are retained in the coastal area (Castro et al. 2000, Castro 2001, Cubillos et al.
2001, Hernández-Miranda et al. 2003). Shelf-break, slope-demersal and mid-water fish species, such
as Chilean hake (Merluccius gayi), big eye flounder (Hippoglossina macrops), and the mesopelagic
Maurolicus parvipinnis, instead seem to prefer early spring reproduction when subsurface waters
drive their eggs and larvae to the coast during the upwelling season, where they develop in the season
of higher production (Vargas et al. 1997, Vargas & Castro 2001, Landaeta & Castro 2002, Landaeta
et al. 2006). This strategy, originally described for some large offshore copepods such as Rhincalanus
nasutus, and more recently observed also in the galatheid Pleuroncodes monodon and the majid
Libidoclaea granaria, is currently accepted as a common feature among several types of organisms
that have in common larvae inhabiting subsurface waters in central Chile (Castro et al. 1993, Yannicelli
et al. 2006a,b). Other species of decapod crustaceans also synchronise their reproduction with the
seasonal changes in hydrodynamics for their transport or retention (Yannicelli et al. 2006b). In this
group, however, several species reproduce during the upwelling season in spring and summer and
their horizontal distribution seems to be associated with their behavioural ability for vertical
migration (i.e., Neotrypaea uncinata, Pagurus sp.). Other species with protracted larval periods
such as Emerita analoga and Blepharipoda spinimana, which reproduce late in summer and early
spring and then reside in the upper water column without signs of vertical migration, probably use
more than a single retention process, as yet unknown, because their reproduction occurs over periods
of contrasting hydrographic processes (Yannicelli et al. 2006b). Among molluscs, there is scarce
information; for instance, the larval retention and shoreward transport mechanism of the gastropod
Concholepas concholepas are the only information reported for the HCS. In this case, avoidance
of the surface Ekman layer by competent larvae appears to be accomplished by an inverse vertical
migration that reduces their time exposed to seaward flow and keeps the larvae between the coast
and an upwelling front (Poulin et al. 2002a,b). Coastal oceanographic processes such as upwelling
shadows have also been reported (Escribano et al. 2002) as larvae retention mechanisms and such
might constitute an understudied coastal larval retention system along the HCS.
Tidal transport of larvae, associated with frontal structures near the coast, the entrance of
estuaries or large bays, has also been reported recently for central and southern Chile. Internal tidal
bores associated with semi-diurnal temperature changes coincided with bivalve, gastropod and
crustacean settlement, suggesting that coastward larval transport occurred during these events in
summer (Vargas et al. 2004). In Corral Bay (~40°S), changes in larval fish distributions coincident
with changes in the estuarine front position in different tidal phases have been reported by Vargas
et al. (2003) as an indication of potential larval tidal transport. More recently, with the aid of fine-
resolution current profiles and stratified larval collections over 24-h cycles, semi-diurnal changes
in water flow patterns and of decapod and larval fish fluxes in and out of the Gulf of Arauco have
been estimated (Yannicelli et al. 2006a, R. Veas unpublished data). The overall larval fluxes were
modified by their vertical position in the water column, the diurnal vertical migration patterns and
the tidal cycle. Interestingly, although clearly associated with tidal phases, the larval fluxes expe-
rienced by these decapod and fish larvae in the Gulf of Arauco did not correspond exactly to
selective tidal stream transport (STST) as it has been reported at the entrance of other bays and
estuaries in other coasts of the world (Forward et al. 1998).
272
Seaward and alongshore exportation of larvae in river plumes, filaments and eddies seems to
be a common feature along the HCS. In northern Chile, a mixture of coastal and offshore larval
fishes, both in the coastal and adjacent offshore areas, has been repeatedly reported in different
seasons and years (Loeb & Rojas 1988, Rojas et al. 2002, Rodríguez-Graña et al. 2005). Besides
the existence of a narrow continental shelf, at least three oceanographic processes involving larval
transport have been advocated to explain such patterns: (1) seaward surface Ekman transport and
upwelling plumes, (2) the presence of filaments, and (3) warm-water intrusion during EN years,
all well-documented processes in northern Chile (González et al. 2000b, Sobarzo & Figueroa 2001).
In central Chile, surface Ekman transport and cold-water filaments have also been reported to export
chl-a, meroplankton and ichthyoplankton from the coastal zone (Cáceres 1992, Morales et al. in
review) and they have been used to explain part of the larval mortality rate estimations in the coastal
zone (Castro & Hernández 2000, Landaeta & Castro 2006). Larval transport associated with river
plumes, mesoscale processes not occurring in northern Chile, has also been proposed as a deter-
minant of barnacle larval transport (Vargas et al. 2006c). Such plumes are also potential areas of
increased larval food availability at the frontal area. Other still-unknown oceanographic processes
capable of transporting larval forms alongshore probably occur in the HCS. In fact, it is worth
noting that almost no reports exist on the role of the Humboldt Current itself or the Chile-Peru
poleward undercurrent as a means of alongshore latitudinal larval transport (except for potential
transport of larval Pleuroncodes monodon in subsurface waters off central Chile; Yannicelli 2005).
In summary, a number of larval transport processes have been described along the central and
southern part of HCS. Three of them seem particularly important when considered from an
ecological perspective: (1) larval transport (seaward and shoreward) in the surface Ekman layer,
(2) exportation from the coast in filaments (especially off the Mejillones Peninsula in northern
Chile and off the Talcahuano area of central Chile) and (3) the coastward subsurface larval transport
in upwelling waters in spring and summer with the concomitant mixing of offshore and coastal
species near shore. Examples of important differences between northern and central Chile are the
influence of oceanographic processes such as EN events (much stronger in northern Chile) and of
river plumes (absent in northern Chile). Overall, populations of invertebrates and fishes along the
HCS develop multiple strategies to cope with the intense periods of transport during early life
stages. Timing the reproductive seasons with specific oceanographic events is most common.
However, specific reproductive timing depends on the local sites of reproduction, capability of
larvae to move vertically in the water column, length of larval life history/cycle and other not so
well studied processes (i.e., retention in upwelling shadows) or those occurring during the adult
life stage (seasonal growth, energy accumulation, oogenesis) that may finally affect the larval
characteristics mentioned above.
Life-history adaptations of macroalgae

The macroalgal flora along the HCS is characterised by the presence of endemic species (32%) mixed
with species of different biogeographical affinities, including subantarctic (34%), widely distributed
(23%), tropical (3.5%) and amphi-equatorial (7%) species (Santelices 1980). In general, this species
composition, which is comparatively poorer than that reported for other regions and increases in
number toward higher latitudes (Lüning 1990, Santelices & Marquet 1998), responds to the relative
biogeographic isolation as a consequence of the predominant direction of the water circulation regime
in the HCS (Santelices et al. 1980, Meneses & Santelices 2000). However, an increasing number of
invasive species as a result of human activities (ship transport, aquaculture, etc.) has recently been
reported with concern about their impact on indigenous species (Castilla et al. 2005a). For example,
recent surveys indicate an increase of subtropical elements, a decrease in endemic species and two
break points in species composition at 12°S and 42°S (Meneses & Santelices 2000).
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MARTIN THIEL ET AL.
Environmental gradients and strategies of resistance, recovery and recolonisation

Although benthic algae are not in direct contact with the main stream of the HCS, subsystems such
as the Arica-Mejillones Current or Gunther Undercurrent are located close to the coast (<100 km)
(Strub et al. 1998) and hence strongly influence macroalgal assemblages. On the other hand, the
biogeographic patterns and ecology of macroalgae of northern-central Chile may be affected by
upwelling events and by changes in the oceanographic conditions due to EN episodes. Both
processes can interact on local and temporal scales: the permanent coastal upwelling can buffer
and moderate superficial warming of the sea and depletion of nutrients during EN (Vega et al.
2005). The first factor has not been well studied for the case of the macroalgae; however, the effects
of EN have received much attention in the last decades (Camus 1990, Vásquez & Vega 2004b).
Over the last 30 yr, studies of seaweed ecology have traditionally focused on conspicuous
genera such as Lessonia, Durvillaea and Macrocystis. Their distribution, reproductive phenology
and recruitment patterns as well as ecological functions on the rocky shores have been well
documented for communities between 40°S and 20°S (Santelices et al. 1980, Moreno & Sutherland
1982, Santelices & Ojeda 1984, Vásquez 1992, Westermeier et al. 1994, Tala et al. 2004 and
references therein). These brown algae represent the major biotic factors structuring the intertidal
and subtidal communities, directly affecting coastal processes through changes in species diversity,
biomass and PP (Ojeda & Santelices 1984, Santelices & Ojeda 1984, Vásquez & Santelices 1990,
Santelices 1991, Vásquez 1992, Camus 1994a, Ortiz 2003, Tala & Edding 2005, Vega et al. 2005).
Because the coastal scenarios along this region are relatively similar in terms of prevailing
oceanographic conditions, it may be argued that most macroalgae are exposed to the same envi-
ronmental factors and thus show similar adaptations. However, considering the phenology, algae
respond to environmental variability basically in two ways (sensu Kain 1989): (1) ‘season antici-
pators’ can grow and reproduce in a strategic annual rhythm suitable for the species, which does
not necessarily require suitable environmental conditions but rather an environmental trigger (e.g.,
day length, nutrient level; Lüning & tom Dieck 1989); and (2) ‘season responders’, by contrast,
grow and reproduce when environmental conditions are favourable (e.g., suitable light conditions).
Whether these two life-strategy mechanisms operate in algae along the HCS has not been well
addressed; however, some patterns of abundance and reproduction as summarised in Figure 21,
based on large geographic distributional scales (described, e.g., for Lessonia, Durvillaea, Macro-
cystis and some genera of Rhodophyta) may be associated with ‘season anticipation’ or ‘season
responses’. However, taking into account that upwelling processes, ENSO or human activities (e.g.,
pollution due to mining discharges) modify local environmental conditions, the existence of other
adaptive mechanisms to withstand these perturbations cannot be ruled out. With the exception of
the well-documented consequences of EN on population dynamics of the dominant kelp Lessonia
(Camus 1994b, Martínez et al. 2003, Vega et al. 2005), questions related to functional strategies of
other macroalgal assemblages during and after mass extinction events within the HCS have been
less addressed. A latitudinal asynchrony in biological processes would be related to different
climatic regimes affecting life-history traits (Mediterranean vs. desert climate in southern-central
and northern Chile, respectively). However, local environmental conditions exert a strong pressure
on the observed patterns and the later recruitment, mainly related to seasonal grazing intensity,
habitat distribution (intertidal versus subtidal) and water movement (Santelices & Ojeda 1984,
Camus 1994a, González et al. 1997, Buschmann et al. 2004b, Tala et al. 2004, Vega et al. 2005).
In this sense, species that reproduce continuously may show advantages in unpredictable environ-
ments compared with strictly seasonal species. Thus, the knowledge about environmental thresholds
for life-history adaptations (expression of functional responses to optimise resource use, e.g., light,
temperature and nutrients, or to withstand environmental stress) is crucial in understanding the
biogeographical patterns of macroalgae.
274
Phaeophyceae Rhodophyta
L.n. (∼20°S)
−20°
C.ch. (∼30°S)
L.t. (∼30°S)
G.ch. (∼30°S)
M.i. (∼30°S)
G.p. (∼30°S) −30°

G.k. (∼30°S)
C.c. (∼30°S)
Intertidal
D.a. (∼32°S) Subtidal
Southern latitude
L.n. (∼32-33°S)
D.a. (∼32-33°S) M.I. (∼35°S)
−40°
M.I. (∼39°S)
L.n. (∼39°S)
D.a. (∼39°S)
M.p. (∼41°S) −50°
100%
50%
Sp Sm A W
Figure 21 Seasonal patterns of reproductive phenology in macroalgae along latitudinal gradients in Chile.
Small inserts show seasonal phenology: Sp, Spring; Sm, Summer; A, Autumn; and W, Winter. Information
was adapted from: Lessonia nigrescens (L.n.) (Camus 1994a, Westermeier et al. 1994); L. trabeculata (L.t.)
(Tala et al. 2004); Macrocystis integrifolia (M.i.) (Buschmann et al. 2004b); Glossophora kunthii (G.k.) (García
1996); Durvillaea antarctica (D.a.) (Santelices et al. 1980, Westermeier et al. 1994, Collantes et al. 2002);
Macrocystis pyrifera (M.p.) (Buschmann et al. 2004b); Chondrus canaliculatus (C.c.) (Vega & Meneses 2001);
Chondracanthus chamissoi (C.ch.) (González et al. 1997); Gelidium chilense (G.ch.) (Dantagnan 1993);
Gastroclonium parvum (G.p) (Rivera 1992); Mazzaella laminarioides (M.l.) (Santelices & Norambuena 1987,
Westermeier et al. 1987).
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MARTIN THIEL ET AL.
Temperature tolerance
Thermal adaptations of the life-history stages, mainly for growth, reproduction and survival require-
ments, are key factors explaining the geographic distribution of benthic algae. In general, two major
aspects characterise temperature demands of macroalgae distributed along the HCS: (1) there is a
close correlation between the upper survival temperature of gametophytes and the northern distri-
bution limits of the species and (2) algae occurring between 18°S and 40°S show higher temperature
tolerance (18–28°C) than algae south of 40°S (17–23°C), coinciding with the latitudinal influence
of the HCS and the boundaries of the two traditionally recognised biogeographic regions (Chilean-
Peruvian and Magellan provinces) (Santelices 1980, Wiencke & tom Dieck 1990, Peters & Breeman
1993, Santelices & Marquet 1998, Martínez 1999). For example, brown algae with Antarctic/
subantarctic distribution commonly found between 18°S and 40°S (e.g., Adenocystis utricularis
and Scytothamnus fasciculatus) have maximal survival temperatures close to 18°C and 24°C
(Wiencke & tom Dieck 1990). These lethal values are above the mean monthly temperatures
measured around 18°S (14–19.5°C) and 37°S (11.7–15.5°C) (Gorshkov 1985). However, when the
upper limit for gametogenesis is regarded, these species do not reproduce at temperatures >13–15°C
(Peters & Breeman 1993). Such thermal requirements suggest that they may survive moderate EN
episodes, but probably a recovery through sexual reproduction may become limited. In the case of
species restricted to the HCS (e.g., the red alga Chondracanthus chamissoi), the growth of both
gametophytes and sporophytes increases at temperatures of 25°C, which is 5–6°C higher than water
temperature in northern Chile (Bulboa & Macchiavello 2001). This is a known phenomenon in
macroalgae and may reflect a reasonable safety limit to survive unpredictable increases of temper-
ature for a long time (e.g., months during EN) or may represent a potential for shifting the
distribution boundaries northward. It must be emphasised that most of the macroalgae from north-
ern-central Chile studied so far show a capability of growth at very low temperatures (>10°C),
indicating clearly an adaptation to the cooling effect of the HCS and reflecting their subantarctic
affinities (Wiencke & tom Dieck 1990, Peters & Breeman 1993, Santelices & Marquet 1998).
Physiological and morphofunctional adaptations
In general, there are only a few ecophysiological studies addressing adaptations of the life history
of macroalgae to varying light and nutrient conditions and they have normally been restricted to
the genera of commercial importance (e.g., Lessonia, Gracilaria, Gelidium, Mazzaella and Por-
phyra) (Oliger & Santelices 1981, Hoffmann & Santelices 1982, Hoffmann et al. 1984, Correa
et al. 1985, Hannach & Santelices 1985, Avila et al. 1986, Bulboa & Macchiavello 2001, Véliz
et al. 2006). Although physiological performances (measured as growth or photosynthesis) are
comparable to those of species from other biogeographical regions, much of the existing information
is site-specific and has been gathered from individual species, indicating adaptations to narrow
ranges of environmental variability. However, in genera such as Gracilaria and Porphyra, which
are exposed to highly changing environmental conditions in enclosed bays, estuaries or upper littoral
zones, broader ranges of environmental tolerance may be expected (Gómez et al. 2004, 2005a).
Due to its physical configuration, the coast along the HCS is characterised by high energy, and
hence physical perturbations such as wave action or sand erosion/accretion fluctuations are impor-
tant and often govern the population dynamics of various infralittoral algae such as Lessonia,
Mazzaella and Gymnogongrus. Thus, algae have developed a suite of morphofunctional adaptations
such as alternation of crustose and erect morphs, large size and seasonal regulation of abundance
(Santelices et al. 1980, Jara & Moreno 1984, Santelices & Ojeda 1984, Santelices & Norambuena
1987, Gómez & Westermeier 1991, Westermeier et al. 1994, Vega & Meneses 2001). In many cases,
these adaptive strategies operate in the early development of the life cycle and in both isomorphic
276
and heteromorphic phase expressions. Processes such as coalescence of spores (Santelices et al.
1999, 2003), regrowth from crusts or vegetative propagation (Hannach & Santelices 1985, Gómez
& Westermeier 1991, Macchiavello et al. 2003), selective mortality of early developmental stages
(Martínez & Santelices 1998), differential phase ratios (Vega & Meneses 2001) and synchronisation
of spore release (Edding et al. 1993, Tala et al. 2004) have been described in algae from northern-
central Chile, which are related to potential selection under changing environmental conditions.
While the seasonal and latitudinal gradients in environmental conditions along the HCS are
recognisable, the magnitude of their impact on the physiological and reproductive biogeography
of benthic algae remains diffuse. Therefore, comparative studies focused on assemblages from
different sites along the HCS are needed in order to define, for example, the environmental
thresholds involved in reproductive fitness, physiological adaptation and recovery capacity.
A special case: enhanced solar radiation
Ozone depletion (with the concomitant increase of UV-B radiation) over the Antarctic region, which
in spring can reach areas as far north as 36°S, has opened the debate about its consequences on
the marine biota of cold and temperate regions (Madronich et al. 1995, Sobolev 2000). Short
wavelengths (UV-B) affect photosynthesis in different ways and have detrimental effects on DNA
and other key cell components (Bischof et al. 2006). Recent studies indicate a potential impact of
current solar UV radiation on photosynthesis of intertidal macroalgae from the southern limit of
HCS (39°S; Gómez et al. 2004, Huovinen et al. 2006). In northern Chile (30°S), zoospores, gameto-
phytes and embryonic sporophytes of subtidal Lessonia trabeculata and the intertidal L. nigrescens
show elevated sensitivity to UV exposure, leading to high spore mortalities and decreases in
germination under current UV doses (Véliz et al. 2006). In general, UV sensibility of early stages
correlates with the depth-distribution patterns of the parental sporophytes, suggesting that this factor
may play a relevant role in depth zonation of benthic algae in this region (Gómez et al. 2005b).
Recent surveys indicate that intertidal species display various photoprotective mechanisms, in
particular the ‘dynamic photoinhibition’, regarded as a down-regulation of the photosynthetic
apparatus to quench the impact of excess energy (Gómez et al. 2004). Some intertidal algae also
have noticeably high contents of UV-absorbing substances (e.g., mycosporine-like amino acids)
(Huovinen et al. 2004). Whether algae from different latitudes exhibit a differential susceptibility
to UV radiation in the context of the HCS remains unclear and future work should give new insights
into the ecophysiological strategies of macroalgal assemblages in scenarios of climate change.
A brief history of exploitation of natural resources in the HCS

First coastal settlers
Evidence of coastline occupation off Peru and Chile may date from as early as 11,000 yr ago
(Llagostera 1979, Muñoz 1982, Báez et al. 1994, Keefer et al. 1998, Sandweiss et al. 1998, Méndez
2002, Méndez & Jackson 2004, Santoro et al. 2005). One of the first studies of the earliest human
habitation along the coast of northern-central Chile (Arica to Coquimbo) was carried out by Junius
Bird in 1941 during excavations of shell middens. This author (Bird 1943, 1946) showed that the
earliest evidence of human settlements on this coast went back to ~6000 yr before present (BP)
and people used highly specialised artifacts with an efficient maritime adaptation. According to
Santoro et al. (2005) evidence of the earliest inhabitants (~11,000 yr BP) is difficult to find since
most prehistoric sites are now on drowned landscapes. However, there is clear evidence that coastal
populations in northern-central Chile used marine natural resources during the Holocene (Llagostera
1979). It is thought that early fishermen dived in subtidal waters to collect molluscs and they also
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MARTIN THIEL ET AL.
speared large fish and marine mammals such as sea lions (Otaria juvata and O. flavescens) (Santoro
et al. 2005). A report on the navigation off the west coast of South America carried out by Lothrop
(1932) showed artifacts used by aboriginal people of northern Chile. Most remarkable was a raft
composed of two cylinders of seal hides tied together to support a small platform on top, and while
voyages out of sight of land were rarely attempted, it was not uncommon to remain at sea for 2 or
3 days (Lothrop 1932).
Llagostera (1979), working between 21°S and 25°S, found evidence of groups of people with
the ability to exploit natural resources from coastal waters ~10,000 yr BP, and he found remains
of molluscan species such as Concholepas concholepas, Fissurella spp., Tegula atra, Choromytilus
chorus, several species of fishes (e.g., Isacia conceptionis and Trachurus murphyi), and semi-
fossilised bones of sea lions and dolphins. Reports from southern Peru suggest specific sites where
mainly fish and seabird resources were exploited (Keefer et al. 1998), while at other (more ephem-
eral) sites they processed mainly molluscs (Sandweiss 2003). Studies from southern Peru and
northern Chile demonstrated that the resources exploited by early coastal settlers of the HCS
changed with time (Llagostera 1979, Sandweiss et al. 2001), which is taken as an indication of
climate change and EN events, causing gradual or abrupt changes in available resources. Similar
observations were made in central Chile near 32°S (Báez et al. 2004). For the late Holocene
(4000–2000 BP) Méndez & Jackson (2004) reported a high degree of mobility of coastal people
in central Chile, who apparently moved between different sites in a region, exploiting the accessible
resources at a given site in an opportunistic manner.
A systematic study of remains of marine invertebrate fauna from central Chile (Curaumilla,
33°S) defined the ecological role of early inhabitants as shellfish gatherers (Jerardino et al. 1992).
According to these authors, they probably modified areas of the rocky intertidal, causing decreases
in mean sizes of Concholepas and Fissurella. Interestingly, Llagostera (1979) suggested that the
appearance of larger shells of Concholepas in shell heaps is indication of an increasing radius of
action and the exploitation of new fishing grounds. This author also remarked that the appearance
of some fish species (e.g., cusk eels, locally called ‘congrio’, from the genus Genypterus, which
can only be fished at greater depths) around 3000 BP is indication that coastal fishermen started
to venture farther out to sea during that time period.
In general, prehistoric people used littoral resources in an opportunistic manner, and during
the past millennia they increasingly widened their radius of action, improved their navigating skills
(rafts) and developed their fishing techniques (fishing nets, hooks). Extraction of marine resources
was not only for subsistence of local groups, but also for an intensive transfer of fish toward inland
sites (Briones et al. 2005). Local people persisted and exploited marine resources until well after
the appearance of the Spanish (Llagostera 1979). The resources collected and captured by prehistoric
people are the same that still today play an important role in the fisheries of northern and central
Chile.
Artisanal benthic fisheries

The present artisanal fisheries in the HCS between 18°S and 35°S are very diverse, comprising
~13 species of algae, ~45 species of invertebrates (molluscs, crustaceans, echinoderms, tunicates)
and ~68 fish species (SERNAPESCA 2005). Before the 1980s, fisheries in general were of low
level and stable since products only went to local markets and the intensity of exploitation was
comparatively moderate (Stotz 1997, González et al. 2006). During the mid-1980s the export of
most of the resources was initiated, producing increases in captures, and once the accumulated
biomass was used up, the resources remained at low and fluctuating levels (Figure 22). In general,
these fluctuations have been interpreted as the classical signs of a badly regulated fishery, as
278
Argopecten purpuratus
5000 25000
Harvest aquaculture (t)

A AMERB
4000 Stock Aquaculture 20000
3000 build-up Fishery 15000 I
18° −
2000 10000
1000 5000
II
0 0
12000 Mesodesma donacium
B III
10000
8000 AMERB
IV
Southern latitude
6000
V 31° −
Landings (t)
4000 RM
2000 VI
0 VII
25000 VIII
C Concholepas concholepas
20000 IX
15000 Closures
10000 Fishery AMERB X 42° −
5000
0 XI
8000
D Loxechinus albus
6000
XII
4000
53° −
2000
0
91
81
01
89
93
83
99
03
79
85
87
95
97
05
19
19
20
19
19
19
19
20
19
19
19
19
19
20
Figure 22 Landings of the four most valuable invertebrate resources in Regions I–VIII, according to fishery
statistics of SERNAPESCA: www.sernapesca.cl and AMERB, ‘Area de Manejo y Explotación de Recursos
Bentónicos’ (MEABR, Management and Exploitation Area for Benthic Resources).
described by Hilborn & Walters (1992): discovery of a stock, development of its fishery and
subsequent overexploitation and eventually collapse.
In order to improve the management status of benthic fisheries (mainly dive fisheries for
invertebrates and algae), Chile has established a system of Territorial User Rights for Fisheries
(TURF), called Areas de Manejo y Explotación de Recursos Bentónicos (AMERB or Management
and Exploitation Areas for Benthic Resources MEABR, which are the diverse names and shortcuts
under which they have been described in the literature) (Castilla 1997, Stotz 1997, Castilla &
Fernández 1998, Aviléz & Jerez 1999, Bernal et al. 1999, Meltzoff et al. 2002, González et al.
2006). This management tool grants exclusive fishing rights over a defined coastal area to legally
established organisations of local fishermen. These areas are exploited according to a management
plan, developed by professionals, approved by authority, and then worked by fishermen under the
permanent supervision of the administrative authority. This system was initiated in practice by
fishermen at the beginning of the 1990s, but legally established in 1997 (Stotz 1997). In general,
it has proven to be a good tool to increase stocks and recover depleted fisheries (Stotz 1997, Castilla
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MARTIN THIEL ET AL.
& Fernández 1998, González et al. 2006), albeit it appears to be deficient in particular situations
(e.g., Gelcich et al. 2006).
Since resource stocks are continuously monitored in management areas, with captures well
controlled and registered, it has been possible to begin to understand the underlying nature of
fluctuating landings. Contrary to the above-described classical pattern of a badly managed fishery,
it has often turned out to reflect the natural variability of the environment of the HCS. Global-scale
phenomena, such as EN events, which produce an outburst of some resources and disappearance
of others, together with more localised processes of upwelling and current systems, generate a
complex, spatially and temporally changing, mosaic of conditions. Fluctuating fisheries are mainly
the consequence of this, the fishermen following (and suffering) natural variations, but not always
causing them, as generally assumed. The following description of the four most valuable resources
fished by artisanal fishermen illustrate this.
Case studies
Scallop fishery
The scallop fishery can be considered as a ‘boom-and-bust’ fishery, where the ‘boom’ is caused
mainly by EN events (Wolff 1987, Stotz 2000, Wolff & Mendo 2000, Stotz & Mendo 2001, von
Brand et al. 2006). During EN, recruitment of Argopecten purpuratus is intense, and during
following years, given the normal fast growth of the species (Stotz & González 1997), huge stocks
of scallops build up, with fluctuations of several orders of magnitude between years (Figure 22A,
stock increased with EN 1982–1983; Figure 23A stock increased with EN 1997–1998) (Stotz &
Mendo 2001). However, following the increased scallop stocks, the development of similar predator
populations and/or the shift of prey preference of predators in response to increased scallop
abundance (Ortiz et al. 2003), together with the fishery, leads to an increasing mortality, which
finally generates the ‘bust’ (Figure 23A) (Wolff 1987, Jesse & Stotz 2002, León & Stotz 2004).
Fishermen are just able to take advantage of part of the EN production before the natural mortality,
caused mainly by predation (e.g., by Octopus mimus), but also by mass strandings (González et al.
2001), takes most of the scallops away (Figure 23B). Normal (LN) years are characterised by small
and, due to spatially (Aguilar & Stotz 2000) and temporally variable recruitment, fluctuating scallop
stocks, which supply a low-level fishery (Figure 22A). Taking advantage mainly of natural recruit-
ment while preventing predation, in northern Chile aquaculture has been able to build up stocks
and harvest at levels several times above fishery production (Figure 22A) (Stotz 2000).
Surf clam fishery
The ‘macha’ Mesodesma donacium fishery also shows boom and bust fisheries along the coast
(Figure 22B). In the past, on most sites a relatively stable low-level fishery existed, produced by
fishermen working in the intertidal zone. During the mid-1970s fishermen learned also to dive for
this resource, which means putting the boat behind the breakers (‘rompiente’), diving, loaded with
at least 40–50 kg of lead, through the surge and then working underneath the breakers. This began
in Region V (32–35°S). After having depleted the local stocks in Region V, some fishermen came
over to Region IV (29–32°S), where the local fishermen quickly learned the same technique. This
has generated a tradition of divers, mainly from Regions IV and V, working on the macha along
the entire Chilean coast, making use of and depleting macha stocks throughout the country (Figure
22A). However, after the establishment of AMERBs for this resource, other reasons for the depletion
became apparent. With the EN 1997–1998 all the macha beds between Arica and Coquimbo,
managed conservatively within AMERBs, died off within a few days. The beds in Coquimbo were
smothered by mud, washed into the bay by a river flood due to heavy rainfall (Miranda 2001). For
280
20
Stock density (ind m−2)

Harvest + A
15 El Niño Predators +
Stranding
10
0
92 93 94 95 96 97 98 99 00 01 02 03 04
12
< 90 mm > 90 mm Harvest
Stock (million ind)
10 B
8
6
4
2
0
n
b
ec
ec
g
ct
ct
Ap
Fe
Ju
Au
Au
O
O
D
D
2000 2001
600 < 60 mm > 60 mm Harvest C

500
400
Stock (t)
300
200
100
0
r
n
g
ct
ec
r
n
g
ct
ec
r
n
Ap
Ap
Ap
Au
Au
Fe
Fe
Ju
Ju
Ju
O
O
D
1999 2000 2001
Figure 23 (A) Stock size of the scallop Argopecten purpuratus in an AMERB, (B) the impact of the harvest
on the decrease of the same scallop stock, and (C) of a Mesodesma donacium stock in another AMERB, ‘Area
de Manejo y Explotación de Recursos Bentónicos’ (MEABR, Management and Exploitation Area for Benthic
Resources).
Arica no specific reason was identified, but increased temperatures have been mentioned as a cause
of mortality for the same species in Peru (Arntz et al. 1987, 1988). In the Coquimbo area, the only
beds left were in Tongoy Bay, which were exploited according to what was considered then a very
conservative strategy. However, despite dynamic (and apparently conservative) management, these
beds also disappeared (Figure 23C) (Aburto & Stotz 2003). Captures were almost irrelevant
compared with the natural decrease (Figure 23C), which was not renewed by recruitment, because
larval supply is spatially (Ortiz & Stotz 1996) and temporally very irregular or absent during many
years. A similar situation was shown for macha beds managed within management areas in Region
VIII, where, due to the almost complete lack of recruitment, the stocks within the AMERBs
collapsed by 2004 (Figure 22B) (Stotz et al. 2004). Following their disappearance during or after
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MARTIN THIEL ET AL.
EN events, the recovery of local macha beds can be extremely slow, as observed in Peru (Arntz
et al. 2006).
The ‘loco’ fishery (sold as ‘Chilean abalone’ in international markets)
The loco fishery makes use of the muricid gastropod, Concholepas concholepas, described as a
top predator in the intertidal systems (Castilla & Paine 1987), but with its fished populations mainly
occurring in the subtidal zone (Stotz 1997). In such areas and given that its main prey items are
suspension feeders (barnacles, tunicates), the species appears more as a browser, through its prey
taking advantage of high PP in the water column near upwelling areas (Stotz 1997, Stotz et al.
2003). Thus, production and consequently fishery landings vary greatly along the coast, with main
landing sites coinciding with the most important upwelling areas (Figure 24) (Stotz 1997). Given
High larval retention with

normally intense recruitment
Moderate larval retention with
temporally variable recruitment 500,000 Harvest AMERB
Harvest (ind)
Poor larval retention with weak 400,000 Pta. Choros
and sporadic recruitment 300,000
200,000
100,000 N° fishermen = 150
0
Upwelling 1999 2000 2001 2002 2003 2004
Chañaral
Loco production
Caldera
Harvest AMERB
25,000 Pto. Oscuro
Harvest (ind)
20,000
15,000
Huasco 10,000
5,000
0
N° fishermen = 30
1999 2000 2001 2002 2003 2004
Pta.Choros
Harvest AMERB
100,000
ague
Harvest (ind)
80,000
Coquimbo 60,000
40,000
20,000 N° fishermen = 48
0
? AMERB 1999 2000 2001 2002 2003 2004
0 200 400 600 800

Los Vilos
Average annual landings (t yr−1)
Figure 24 Concholepas concholepas: Variability of larval retention and recruitment along the coast of Regions
III and IV, production (average of the period 1985–1995) along the coast of Region IV and harvests of three
AMERBs located at coastal areas differing in production (note the scale of y-axis). (Figure adapted from
J. González et al. (2004) and Stotz (1997)) Harvest data of AMERB obtained from SERNAPESCA (www.
sernapesca.cl); stars show for Region IV the approximate locations of the AMERBs, ‘Area de Manejo y
Explotación de Recursos Bentónicos’ (MEABR, Management and Exploitation Area for Benthic Resources).
282
its complex reproductive biology, which starts with reproductive aggregations for copula, the laying
of egg capsules in which the larvae develop for about 30 days, then followed by a 3-month period
of pelagic life, the result is high recruitment variability at temporal and spatial scales, depending
on coastal oceanography and topography, and the potential for retention areas (Stotz 1997,
J. González et al. 2004) (Figure 24). Larvae settle and metamorphose mainly on adult barnacles
covering adult shells (Manríquez et al. 2004) or in association with recently settled barnacles (Stotz
et al. 1991b), which may vary greatly between years and sites (Stotz et al. 1991a). This produces
a very complex metapopulation structure (J. González et al. 2004, 2006). However, given its long
life, with individual growth varying greatly (depending on food availability), fluctuations become
partly attenuated when the individuals finally recruit to fisheries at an age between 3 and 4 yr with
a size of 10 cm of peristomal opening length (Pérez & Stotz 1992, Stotz & Pérez 1992, Stotz 1997).
Thus, while spatial variability of the fisheries is great, the temporal variability at each site becomes
partially attenuated (Figure 22C after 1996), with fluctuations at the level of two to three times,
which nevertheless means significant changes in the income for fishermen (Figure 24, harvest in
AMERBs). The establishment of AMERBs, which was mainly motivated by the closure of the loco
fishery at the same time (1989–1992), coincided with a period of increased loco stocks along most
parts of the central Chilean coast, probably favoured by the closure and a short period in which
fishermen agreed to strictly obey that measure (Stotz 1997). The relationship of the number of
fishermen to the size of the management area was at that time generating an acceptable income,
but this situation has mostly changed drastically in the following years. At present, many fishermen
located at coastal areas with low production are dissatisfied and willing to abandon their AMERBs
(compare magnitudes of harvest of AMERBs and number of fishermen in the organisation, shown
in Figure 24).
Sea urchin fishery
The urchin Loxechinus albus fishery in the HCS between 18°S and 35°S is perhaps one of the most
variable ones, in this case natural variability probably being increased as a consequence of captures.
In central Chile (Regions IV–VIII), urchins are restricted to shallow areas with great surge on the
exposed coast, in which the species is partly safe from the predation of the rock shrimp Rhyncho-
cinetes typus (Stotz 2004), a species with abundance and distribution pattern that responds to the
variable existence of refuges from its own predators (Caillaux & Stotz 2003). This produces a very
patchy distribution of sea urchins. Recruitment occurs mainly inside the adult aggregations, where
recruits are protected by the spine canopy (Stotz et al. 1992). Thus, when fishermen, taking
advantage of calm weather conditions, reduce the stocks, subsequent recruitment is probably heavily
affected. The observation is that once a site is fished for this species, it only recovers about 10 yr
later (Stotz 2004). Further north (Regions I–III, mainly between 20°S and 23°S), the fluctuations
attenuate slightly and landings increase, as large Macrocystis beds appear (Stotz 2004). There, a
more conservative exploitation is carried out inside the AMERBs, conserving patches, and this
strategy has allowed an increase in numbers, the sustainable exploitation of which nevertheless still
needs to be demonstrated (Figure 22D).
Resource dynamics and management of artisanal benthic fisheries

The great natural spatial and temporal variability of resources, and hence fishery production, which
characterises the HCS between 18°S and 35°S, poses a great challenge to management. Fishermen,
through AMERBs and legal restrictions, are not allowed to move away when a resource goes
through a low cycle, as they used to do in the past. Illegal movement, though still occurring, is
also increasingly prevented in practice by conflicts with the respective local fishers. This produces
283
MARTIN THIEL ET AL.
the risk that fishermen continue fishing on a weakened population, perhaps producing its collapse
to a degree where recovery is severely compromised. Additionally, the distribution of fishermen
along the coast within each region in general is not well adjusted to its spatial productive variability,
thus creating great differences of income along the coast, with fishermen’s organisations located
at productive sites getting increasingly richer, and others at unproductive sites (as shown with
harvest in AMERBs in Figure 24) getting increasingly poorer, which could be a source of conflicts
(Gelcich et al. 2005, Stotz & Aburto 2006). The challenge is to advance to an integral management
strategy in which fishermen, instead of rotating between fishing zones along the coast, rotate
between fisheries of different resources or among other related activities (processing, tourism, etc.)
in order to produce income during years or months of poor production. This means a change from
a specialist to a generalist strategy, but with very strict control of their numbers, such that they are
well adjusted to local production levels. Complementary stock enhancement, using biological and
ecological knowledge and aquaculture experience, might help to mitigate natural fluctuations (Stotz
et al. 1992, Zamora & Stotz 1994, Pacheco & Stotz 2006). In order to avoid the risk of increased
fishing pressure (caused by restricting movements of fishermen) on already weakened populations,
the establishment of a network of small reserves along the coast should be considered, which should
also aid in reducing natural recruitment variability (see also Stotz & Aburto 2006).
Bioeconomic aspects of industrial crustacean fisheries

An important crustacean bottom-trawl fishery exists in northern and central Chile, particularly
between Regions II and IV. This fishery, which includes the nylon shrimp (Heterocarpus reedi),
yellow squat lobster (Cervimunida johni) and the red squat lobster (Pleuroncodes monodon), is
conducted on the continental shelf at depths ranging from ~100 to 600 m. The exploitation of these
resources began in the 1950s as fauna accompanying catches of hake Merluccius gayi and was
subsequently developed into a fishery specific for these crustaceans (Arana & Nakanishi 1971,
SUBPESCA 1999a,b). A growth phase for these fisheries occurred between 1958 and 1968, during
which annual landings of nylon shrimp reached 11,000 t. Subsequently, landings declined to less
than 3000 t in 1979. Between 1980 and 1986 landings remained below 4000 t, except for 1983,
when landings exceeded 6000 t. The last period is interesting in its analysis since it coincided with
three fundamental factors: (1) from 1979 to 1982 the exchange rate (Chilean peso/U.S. dollar) was
39 Chilean pesos/U.S. dollar, which produced a depressive economic effect, resulting in the collapse
of fisheries companies; (2) in 1983 the occupation of new fishing grounds south of Region IV
increased the catch levels above those previously obtained; and (3) from 1984 to 1986 a reorgan-
isation of the fishing fleet occurred due to the opening of the fishery for the red squat lobster
(Pleuroncodes monodon) along with an increase in world prices for this resource. Between 1989
and 1991 the fishery was closed in Regions V and VIII, but it remained open in northern Chile. In
this northern area after 1986 there was an increase in landings of squat lobster reaching 10,620 t
in 1995. After this year there was a clear decrease in these landings, reaching 4000 t in 2000.
Since 1995, the three species are subject to catch quotas per boat owner, based on a total
allowable catch (TAC), which is determined annually, following direct resource evaluations. A
period of difficulty in the crustacean fisheries due to decreasing stocks began in 1999 in northern-
central Chile (SUBPESCA 2005a,b). The TAC per Fisheries Unit of nylon shrimp (Regions II–VIII)
dropped over four subsequent years from 10,000 t in 1997 to only 4000 t in 2000 (i.e., a 60%
decline). The TAC of the fishery for the yellow squat lobster in Regions III and IV dropped from
6000 t in 1998 to 4000 t in 2000, representing a reduction of 33% during that time period. This
substantial reduction in landings caused a major decline in the activities of the fishing fleet and the
packing plants. During 2002, some companies closed, with important losses in employment
producing socioeconomic impacts in the regions affected (Pérez 2003, 2005). This brief overview
284
underscores that the fishery of these benthic crustaceans is highly dynamic, driven not only by
biological factors, but also by administrative decisions and economic considerations. In the follow-
ing section some relevant information on the biology of these three species is provided before
presenting a case study highlighting the relationship between catch-based stock estimates and
species biology.
Basic biology of nylon shrimp and squat lobsters

The main bathymetric distribution of these species shows a strong overlap with the OMZ between
~50 and ~600 m, where they occur on gravel and mud bottoms (SUBPESCA 1999a,b). Their
latitudinal distribution ranges from 25°S to 39°S (Heterocarpus reedi), from 6°S to 40°S (Pleu-
roncodes monodon), and from 29°S to 38°S (Cervimunida johni) (Acuña et al. 1997, 1998, Quiroz
et al. 2005). Little is known about their food resources. All three species themselves are important
prey organisms for demersal fish predators (e.g., flounders and hake), and it has been discussed
that the OMZ may represent a refuge from predation (Villarroel et al. 2001). Most information
about the biology of these crustacean species is based on analysis of specimens obtained from
commercial or research catches.
Heterocarpus reedi
Available data suggest seasonal migrations with the main concentrations of shrimp found at great
depths (>400 m) in austral summer, at shallow depths during austral winter (<200 m), and at
intermediate depths in autumn and spring (SUBPESCA 1999a). During austral summer/early
autumn, a comparatively large proportion of individuals has a soft carapace, which indicates that
moulting is frequent during this time period (SUBPESCA 1999a), possibly accompanied by mating.
Following this period, ovigerous females are found throughout winter and early spring
(May–November) with a peak abundance in July–August (SUBPESCA 1999a). Larval release is
initiated in September. Fecundity has been estimated at 1000–10,000 eggs, and it is generally
assumed that each female produces one clutch per year (SUBPESCA 1999a). Catches have con-
sistently shown higher proportions of females than males, with males smaller in size than females.
Cervimunida johni
Moulting and copulation are thought to occur mainly during the summer months (SUBPESCA
1999b). Embryo incubation extends from May to November, with the highest proportion of oviger-
ous females observed in July–August (in September–October in the northern zone) (SUBPESCA
1999a,b). Larvae are released during the spring months. Fecundity has been estimated at 500–14,000
eggs in Regions III and IV, although some females in these and other regions have been found with
>20,000 embryos. Males reach substantially greater sizes and weights than females, and average
size at first maturity (of females) varies substantially among years (Acuña et al. 2005). It is generally
assumed that each female produces only one clutch per year, and Arancibia et al. (2005) suggested
that C. johni is a slow-growing species, with individuals living up to 11 yr.
Pleuroncodes monodon
Based on the main occurrence of ovigerous females it has been suggested that mating occurs mainly
in late summer/early autumn (Palma & Arana 1997). Embryo incubation extends from April to
November, with the highest proportion of ovigerous females observed in July–August (Palma &
Arana 1997). All embryonic developmental stages were found between June and October (Palma
& Arana 1997), suggesting that not all females mate at the same time. The first planktonic larvae
285
MARTIN THIEL ET AL.
appeared in June, and early developmental stages were found in the plankton until December (Palma
1994), supporting the suggestion that reproduction is not synchronised among females. Gallardo
et al. (1994) reported the first benthic stages in March with very high densities of recently settled
squat lobsters in April. Roa et al. (1995) identified nursery areas in places with very low oxygen
concentrations from where juveniles migrate to nearby adult habitats. It is generally assumed that
females produce only one clutch per year, but the extended reproductive period and the apparently
short incubation period of 2–3 months (Palma & Arana 1997) could allow some females to produce
more than one clutch per year. Fecundity has been estimated at 1800–34,000 eggs in Region VIII
(Palma & Arana 1997), and most females reach sexual maturity at carapace lengths of ~25 mm.
Red squat lobsters are estimated to live from 5 to 10 yr (Roa & Tapia 1998).
Biology and stock estimates

As can be seen from the preceding section, information on the biology of the nylon shrimp and
the squat lobsters is mainly based on demographic data (size at first reproduction, fecundity, sex
ratio, per cent ovigerous females, embryo developmental stages), but little information is available
on their behaviour. A close analysis of monthly landings suggests possible seasonal changes in
behaviour during the year and indicates that it is important to incorporate this information in stock
estimates (Pérez 2005).
Pérez (2005) proposed that the biomass of the nylon shrimp showed well-defined cycles of
availability (sensu Menge 1972), which were not related to its true abundance in biomass (sensu
Menge 1972). The first cycle (termed the ‘short cycle’) occurs from September to December and
is characterised by a decrease in availability (minimum in October), followed by a recovery
(maximum availability in December). This leads to a cycle of greater length (termed the ‘long
cycle’), characterised by a decline in available biomass, reaching a minimum in April and followed
by an increase again reaching a maximum in August. Reasons for these cycles are not clear, but it has
been suggested that they are related to the moult and reproduction. In this sense, a sudden decrease
in availability could be due to recently moulted individuals hiding from predators (Pérez 2005).
The total biomass of Heterocarpus reedi in Region IV in September 1997 was estimated to be
about 5000 t, but not all of these shrimp were susceptible to fishing gear, resulting in an available
biomass (to the fishery) of 4200 t (for details see Pérez 2005). The estimated total biomass decreased
to 2000 t in August 2000 (Week 143), representing a decline of 60% from that estimated at the
initiation of the simulation (Figure 25A). However, the percentage variation in the catch per unit
effort (CPUE) was not proportional to the decrease in abundance within periods of maximum
availability (Figure 25A). At the beginning of the study period, the CPUE was 0.59 t haul−1, and
in August 2000 this decreased to 0.45 t haul−1, representing a decrease of only 24%. In recruitment,
at each fishing station in 1997–1998 and 1998–1999 the difference between the control curves and
availability allowed the identification of two time windows for recruitment, one of lesser intensity
in December and one of greater intensity extending through June and July. This seasonal variation in
the availability of the resource to the fishing gear has a direct influence on the population estimates
and consequently the determination of TACs. Therefore it was recommended to include this biolog-
ical information in stock evaluation and to conduct the corresponding research cruises during periods
of maximum availability of the resource (Pérez 2005).
Crustacean fisheries and economic considerations

Fisheries respond not only to fluctuations in stock abundance but also to economic considerations.
From the economic perspective, the annual TAC system has produced an increase in the fishing
286
1997 1998 1999 2000

6000
5000
Biomass (t) Available biomass
4000
Control biomass
3000
2000
1000
0
0 20 40 60 80 100 120 140 160
Week
A
Income Costs
20 7
Million US$
16 6
5
12 4 Cervimunida johni
8 3 Heterocarpus reedi
2
4 1
0 0
1997/98 1998/99 1999/00 1997/98 1998/99 1999/00
Fishing season
B
Figure 25 (A) Biomass dynamics of the nylon shrimp in Region IV. Available biomass is the biomass that
is available to the fishery and control biomass is the estimated total biomass of the resource according to the
model by Pérez (2005); bars above figure represent short cycles (light shading) and long cycles (dark shading)
when resource becomes unavailable to fishery for biological reasons; for further details see text and reference.
Weeks are numbered beginning with the week of 1 September 1997. (B) Comparison of total income with
total costs by resource of crustacean trawl fishery in Region IV.
effort, measured in terms of hauls, resulting in a decrease in the expected economic benefit based
on administrative measures. Pérez (2003) explored the bioeconomic effect produced by the decrease
in CPUE in the nylon shrimp and yellow squat lobster fisheries in northern Chile during 1997–2000.
A biological-technological simulation model was used by Pérez (2005), which took both physical
and biological variables into account. An economic submodel was incorporated into this model in
order to carry out an integrated analysis of the crustacean fisheries, which included the subsector
involving catches and their processing. The economic results obtained, when integrated with the
catch results and size of the stocks, allowed the dynamics of this fishery to be explained in
Regions III and IV during 1997–2000, when the catch increased by 21%, and final production
increased by 24% (measured as frozen tails), although the variable costs increased by only 11%
(Figure 25B). In this same period the fisheries costs increased by 117% and the total production
cost increased by 93%. The results showed that part of the economic benefit was lost due to the
effect of a decrease in the biomasses of both resources and an excessive increase in the average
production costs (due to increased costs of fishing and extraction).
This example underscores the importance of incorporating economic reference points in addi-
tion to biological (biomass) and fishery variables (CPUE and catch). Furthermore, basic biological
data (larval ecology, settlement biology, habitat requirements, seasonal migrations, and mating
behaviour) still need to be revealed for these three crustacean species. At present, very little is
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MARTIN THIEL ET AL.
known about the influence of oceanographic factors on recruitment success and stock dynamics of
crustaceans from the continental shelf off northern-central Chile. Most current information suggests
that their life history is driven by seasonal factors, and presently available data do not permit an
examination of the effects of interannual variability in oceanographic conditions (e.g., ENSO) on
their population dynamics. Future studies should address these questions in order to achieve a
sustainable fishery.
Pelagic fisheries and fisheries management, 1980–2005

The Chilean purse seine fleet mainly exploits five pelagic fish species: anchovy (Engraulis ringens),
jack mackerel (Trachurus murphyi), chub mackerel (Scomber japonicus), and Pacific sardine or
pilchard (Sardinops sagax) and common sardine (Strangomera bentincki). There were doubts about
the taxonomic status of jack mackerel (Stepien & Rosenblatt 1996, Oyarzún 1998), but a recent
molecular study revealed that the name Trachurus murphyi should be conserved (Poulin et al. 2004).
In the official landing statistics it is also recorded as T. murphyi, but the Technical Reports of the
Undersecretary of Fisheries refer to it as T. symmetricus.
History of the catches

Aguilar et al. (2000) described the development of the Chilean pelagic fisheries based on the
landings of anchovy, jack mackerel and pilchard from 1970 to 1995. Herein the period from 1995
to 2005 (also known as the ‘regulated’ period) is included, which adds two additional species: chub
mackerel and common sardine, and their landings since 1980.
The total annual landings captured by the Chilean purse seine fleet during the study period showed
a steadily increasing tendency from 1980 until 1995, from 3.4 million t to almost 6.9 million t,
driven mainly by the increase in jack mackerel landings and secondarily by anchovy landings,
which replaced the Pacific sardine, the most important species during the early 1980s (Figure 26).
After that, the total annual catch decreased to around the same level found at the beginning of the
study period, with the exception of 1998, when both species simultaneously showed a sharp decline
in their landings. However, as stated, these are ‘regulated’ landings since catches are the result of
fixed total annual quotas and therefore not necessarily representative of resource availability.
Administratively, four of the five species included in this analysis are latitudinally assigned
to fisheries units, each of which comprises two to five administrative zones: Regions I–II
(18°25′S–26°06′S), Regions III–IV (26°06′S–32°18′S), Regions V–IX (32°18′S–39°37′S) and
Regions X–XII (39°37′S–56°30′S), and there is a strong difference in landings between the respec-
tive fishery units (Figure 26). Therefore, annual landings are analysed by species and these latitu-
dinal fisheries units to visualise their importance in each of them. In the following text each Fisheries
Unit is defined by the Regions it contains, e.g., Fisheries Unit I-II.
Landing statistics of anchovy show the typical characteristics of engraulids, with successive
increases and decreases, attaining highest landings of 2.7 million t in 1994, but landings usually
do not surpass 1.5 million t. Anchovy landings are most important in northern Chile (Fisheries
Unit I–II) (Figure 27). In contrast to all other species, jack mackerel showed a continuous increase
until 1995, when annual landings reached 4.4 million t, after which catches continuously decreased
again. Throughout this period, catches in central-southern Chile (Fisheries Unit V–IX) had over-
riding importance (Figure 27). The highest landings of chub mackerel were attained in 2003 with
0.5 million t. This species has shown a steady increase in landings during the study period. At the
beginning of the period, landings of chub mackerel were mainly in northern Chile (Fisheries
Unit I–II) but since the year 2000 also increased in central-southern Chile (Fisheries Unit V–IX)
288
Figure 26 Total annual landings for the five most important pelagic species caught by the Chilean purse seine
fleet during the period 1980–2005; left panel shows SST anomaly, total landings and landings of the five
species; right panel shows average landings per latitude and year in each of the four fisheries units, calculated
for the time period 1980–2005.
(Figure 27). The highest landings of Pacific sardine were attained in 1985 with almost 2.9 million t.
During the 1980s this species was the most important small pelagic fish captured in Chilean waters
but catches continuously decreased during the late 1980s, reaching very low levels in the mid-
1990s. The Pacific sardine was most important in northern Chile (Fisheries Unit I–II) but gained
proportionally in importance in central-southern Chile (Fisheries Unit V–IX) during the early 1990s
(Figure 27). The highest landings of common sardine were attained in 1999 with 0.75 million t.
During the 1980s this species was very scarcely captured in Chilean waters but from then on showed
a steady increase (Figure 27). Landings of the common sardine were most important in the central-
southern Chile (Fisheries Unit V–IX).
Relationships with oceanographic variations

The coastal areas off the Chilean coast are known for being typical of an EBC system, where
upwelling is a characteristic oceanographic feature. Fonseca & Farías (1987) and other authors
described the presence of active upwelling centres in several areas of the Chilean coast, like Iquique
(Fuenzalida 1990) and Antofagasta in Fisheries Unit I–II (Blanco et al. 2001), Caldera and
Coquimbo (Acuña et al. 1989) in Fisheries Unit III–IV, Valparaíso (Johnson et al. 1980) and
Concepción (Cáceres & Arcos 1991) in Fisheries Unit V–IX.
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MARTIN THIEL ET AL.
Niño 3.4 index

2
1
0
−1
−2
−3
2.500
ANCHOVY
2.000
1.500
1.000
500
0
4.500
JACK MACKEREL
3.500
2.500
Landings (1000 t)
1.500
500
0
350
CHUB MACKEREL
250
150
50
0
3.000
PACIFIC SARDINE
2.000
1.000
0
800
COMMON SARDINE
600
400
200
0
1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004
Figure 27 Total annual landings for the five most important pelagic species caught by the Chilean purse seine
fleet during the time period 1980–2005 in the respective fisheries units; grey dots represent Fisheries Unit I–II,
open dots Fisheries Unit III–IV, grey triangles Fisheries Unit V–IX, and open triangles Fisheries Unit X–XII.
Yáñez et al. (1996) conducted a survey to assess the possibility of introducing the use of SST,
obtained from NOAA (National Oceanic & Atmospheric Administration) satellite data, for the small
pelagic fisheries resources and found significant relationships between species yields and TGRs.
Jack mackerel yields were largely related to strong TGRs next to oceanic waters, while anchovy
and common sardine yields were mainly associated with the development of coastal upwelling
events. Comparison with the SST anomalies shows that landings of anchovy negatively correlate
290
with SST anomalies (Yáñez et al. 2001); also chub mackerel landings seem to correlate with SST
(Figure 27). In contrast, interannual variations in the landings of the other three species seem to
be largely independent of variations in SST (Figure 27). The stabilisation of maximum anchovy
landings between 1.5 and 2.7 million t during the 1990s and the parallel decline of the landings
of the Pacific sardine in the HCS (and the entire southeastern Pacific) reflects another regime shift
from the warm ‘sardine regime’ to a cool ‘anchovy regime’ (Chavez et al. 2003, Alheit & Niquen
2004, Halpin et al. 2004). These regime shifts occur on multidecadal scales and are probably related
to the PDO, but the mechanisms driving these changes are not yet well understood.
Silva et al. (2003) studied the relationship between chl-a concentration, SST, and fishing yields
of anchovy, Pacific sardine and jack mackerel in northern Chile during summer and autumn 1999.
CPUE superimposed over SST images confirmed that the anchovy has a more coastal distribution
than Pacific sardine and jack mackerel, being found in the frontal zone of coastal areas. In the
southeastern Pacific, the jack mackerel is a heavily exploited pelagic species, and its presence in
the HCS in autumn and winter is assumed to be mainly due to an inshore feeding migration
(Bertrand et al. 2004). During warmer years, jack mackerel may immigrate into coastal waters
where they are thought to exert high predation pressure on anchovy (Alheit & Niquen 2004).
Changes in SST associated with EN events may also affect the migration pattern of jack mackerel,
which needs to be taken into account in stock assessment and management plans (Arcos et al. 2001).
Management of pelagic fisheries

According to Aguilar et al. (2000) the traditional method used to conserve fish stocks and prevent
overfishing is to set a TAC for the fishery. Typically, TACs aim to restrict fishing effort to its MSY
(maximum sustainable yield) level. Once these ‘safe biological limits’ are reached, fishing is
prohibited. But TACs do not, by themselves, address the overcapitalisation issue. Consequently,
many fisheries economists recommend that the designated TAC is distributed to industry participants
in the form of individual transferable quotas (ITQs), quasi property rights that restrict additional
access to the fishery. Under these rules, failure to acquire an ITQ effectively forces vessels out of
the fishery, thereby reducing fishing effort and increasing harvesting efficiency.
The Chilean General Law of Fisheries and Aquaculture considers three types of access to the
fishery: (1) Full Exploitation Regime, which includes setting an annual quota or TAC by fishing
unit, which can be temporally divided during the year and also modified once during that same
period (once the full exploitation regime is assigned to a given species no more fishing vessels are
allowed to enter the fishery); (2) Recovering Fishery Regime is the fishery under a state of
overexploitation, subject to a capture ban of at least 3 yr, to obtain its recovery and where an annual
quota (or TAC) can be established; and (3) Early Developing Fishery Regime, which is a demersal
or benthonic fishery with open access where an annual quota can be established and where no
fishing effort is applied or if it is done it is less than 10% of this quota.
In 1993 the first pelagic fisheries in Fisheries Unit I–II were assigned the status of Full
Exploitation Regime, and by 2000 this had extended to four of the five fisheries (anchovy, jack
mackerel, Pacific sardine and common sardine) and to all fisheries units. The chub mackerel is one
of the few commercial species that is still under ‘open access’, with no regulation to date.
Alternative management tools have been developed and used in the last years: in 1998, the use
of a geographical positioning system for the industrial and artisanal fleets was established; later in
2001 the Límite Máximo de Captura por Armador (LMCA, maximum capture limit per owner)
was introduced for the industrial fleet in the fisheries for the small pelagics anchovy, sardines and
jack mackerel in all fishery units, which is essentially an ITQ, and at least had the effect of reducing
the fishing fleet. The LMCAs are determined using captures (1997–2001) and corrected hold
capacity (authorised hold capacity in cubic metres times authorised area length divided by total
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MARTIN THIEL ET AL.
length of fishery unit, Law 19.173). This action was set to last until 2002, but was later extended
until 2012 (Law 19.849). Finally, in 2004 a similar management tool was established for the artisanal
fleet, the Regimen Artesanal de Extracción (RAE, Artisanal Capture Regime), which in this case
assigns the artisanal fraction of the Annual Global Quota to one or more artisanal fishermen’s
organisations in each fishery unit, which in turn divide it between members.
As becomes evident from the preceding paragraphs the currently valid management rules (full
exploitation, Global Annual Quota, LMCA, etc.) have been implemented when landings started to
decrease or stocks had already dramatically declined. Therefore, fisheries of small pelagic fisheries
in the HCS off Chile are presently managed at much lower population levels than they used to be
in previous decades. In fact, stocks of anchovy and common sardine in central Chile are considered
to be overfished (Cubillos et al. 2002). The interaction between administrative effects on landings
and fishery-induced impacts on stocks complicate the detection of direct relationships between
environmental factors and fish stocks. In addition to fishery-independent stock surveys, studies of
the basic biology of individual species, of biological interactions (predators, competitors, food),
and of the role of climatic and coastal oceanography are required in order to improve our under-
standing of the factors driving the population dynamics and distribution in northern-central Chile.
Aquaculture
According to the latest reports published by the Food and Agriculture Organization of the United
Nations (FAO 2006), the contribution of aquaculture to the world supply of fish and shellfish
“continues to grow faster than any other productive sector of animal food origin”. The world
production of aquaculture registered in 2002 rose to 51.4 million t (including aquatic plants), with
Asian countries producing 91.2% of this. In 2002, Chile contributed only 1.4% of the world’s total
production, but it is among the 10 countries showing the fastest growth in aquaculture production.
Fisheries and aquaculture are for Chile one of the most important economic activities with a total
income of U.S. $2,246 million in 2003, of which aquaculture contributed U.S. $1600 million. The
largest share of the Chilean aquaculture production (80%) is from southern Chile (41–46°S), with
salmon and mussels and to a lesser extent oysters, seaweeds and more recently red abalones (Haliotis
rufescens) being the most important resources (FAO 2006).
Aquaculture activities along the coast of northern-central Chile (18–35°S), although not reach-
ing the same levels as in southern Chile, have been continuously growing during the past two
decades (FAO 2006). Given that the shorelines of northern-central Chile are relatively exposed to
wave action and fully subjected to the effects of ENSO, it is particularly important to take these
factors and interannual variability in oceanographic conditions into account. In fact, all aquaculture
centres in northern and central Chile are located in relatively sheltered bays. The main natural
resources cultured in northern-central Chile are scallops (Argopecten purpuratus) and seaweeds
(Gracilaria chilensis), and their natural populations are also exposed to strong seasonal and inter-
annual variations. Small-scale culture of some other species (bivalves Mesodesma donacium and
Tagelus dombeii, gastropods Concholepas concholepas and Fissurella spp., sea urchins Loxechinus
albus) has also been attempted but has not reached a commercial stage, mainly due to biological
(long larval periods) and logistic (food supply) reasons. Several introduced species are also cultured
in northern Chile, namely the Pacific oyster (Crassostrea gigas), which has been cultured on a
small scale since 1970, and during recent years increasingly abalones (Haliotis rufescens and
H. discus hannai).
The main resource cultured during the past two decades in northern Chile is the scallop
Argopecten purpuratus. Culture centres are located in bays, namely Isla Santa Maria and Bahía
Mejillones (22°S), Bahías Caldera, Calderilla and Inglesa (27°S), and Bahías Guanaqueros and
Tongoy (30°S). Standing stocks and productivity in these bays are below those of similar bays in
292
Peru (Uribe & Blanco 2001). It has long been recognised that natural scallop stocks vary together
with ENSO variations, mainly because settlement of small settlers (spat) is strongly favoured during
EN events (Narvarte et al. 2001). Due to this relationship, the local scallop industry is affected in
a positive way by EN, which can lead to an increase in spat collection by >300% (Illanes et al.
1985). In a research project on the effect of environmental factors on scallop culture, data of water
temperature, gonad indices, larval abundance and recruitment (spat collection) were gathered
between 1981 and 1984 in Tongoy Bay (Illanes et al. 1985). The EN event led to a temperature
increase of 2°C above the pre- or post-EN levels at the sea surface and of 2.5°C at the bottom
(20 m). During the EN period, the Gonad Index of adults registered a maximum (25) and a minimum
(6.8) with a massive evacuation of gametes, a situation never observed during normal years (Illanes
et al. 1985). Similar observations were made in southern Peru by Wolff (1988), who concluded
that A. purpuratus is a continuous spawner with spawning peaks during late austral summer and
autumn (February–March). Under the unusually high water temperatures during EN 1982–1983
(2–2.5°C above the normal temperatures), the recuperation time between two spawnings was
shortened, indicating that maturation was accelerated and spawning probably intensified under these
conditions. This interpretation was confirmed by the highest larval concentration found within the
period. Wolff (1988) stated that it is not clear along the Peruvian coast that past EN favoured the
scallops stocks. The EN 1972 apparently did favour stocks, as catches were significantly higher than
during the 3 yr before and after this event, but the weaker EN 1976 did not have the same effect.
In 1979, which was a ‘normal’ year, the catches still exceeded the catches of 1972. Limo (in a
personal communication to M. Wolff) reported enormous numbers of A. purpuratus during the
strong EN 1925 in Ancon Bay, north of Lima. In order to reduce the dependency on natural variations
in supply of small recruits, intense efforts have been undertaken to produce settlers in the laboratory
(Uriarte et al. 2001), but this only satisfies part of the requirements for scallop spat. The fact that
natural spat is still obtained at much lower costs than spat produced in the laboratory may have
led to the strong increase in spat collectors observed during recent years in Tongoy Bay (Figure 28).
Possibly, the decreasing spat collection efficiency between 1998 and 2001 (when the total number
of spat collectors in the bay exceeded 2.5 million bags) is indication that the carrying capacity is
reached and that spat collectors are starting to compete for the available settlers.
Culture of Gracilaria chilensis in northern Chile is mainly developed on shallow soft bottoms
in sheltered parts of the bays (e.g., Pizarro & Santelices 1993). Since sheltered bays are relatively
scarce along the cost of northern and central Chile, aquaculture of this seaweed in this area does
not reach the amounts harvested in southern Chile. Edding & Blanco (2001) observed a decrease
in productivity and yield of ‘agar’ from G. chilensis cultured in Region IV (29°59′S) during EN
1997–1998. This may be more related to the decrease in nutrient concentrations and increase of
visibility instead of the higher water temperatures. Furthermore, these authors cited González
(1998), who reported that increased wave action during EN resulted in an important reduction of
biomass. Growers of G. chilensis in Region IV also reported heavy damage to culture fields on
shallow subtidal soft bottoms caused by storms during EN (R. Rojas personal communication).
However, Santelices et al. (1984) stated that recovery of Gracilaria sp. after storms is unexpectedly
fast due to regrowth of thalli from the portions buried in the sand. Overall, the production of
G. chilensis in northern Chile does not seem to be severely affected by ENSO, but is rather
determined by stock density and harvesting frequency (Pizarro & Santelices 1993). The relatively
limited interannual variability in extracted and cultured biomass (Figure 28) further supports the
suggestion that management rather than environmental conditions drive the production of G. chilensis
in northern Chile.
During recent years first attempts have been made to culture large kelps from northern and
central Chile (e.g., Edding et al. 1990, Edding & Tala 2003) in order to satisfy the growing needs
of the abalone culture. This is considered particularly important since EN can have dramatic impacts
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MARTIN THIEL ET AL.
Niño 3.4 index

3
2 A
1
0
−1
−2
−3
1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006
600 30
B
500 25
Collectors (100,000 bags)

N° of scallop seeds by collector bag
Scallop seeds (n bag−1)
N° of collector bags
400 20
300 15
200 10
100 5
0 0
1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006
20
C
16 Natural Gacilaria production
Harvest (1000 t)
Cultivated Gacilaria production

12
0
1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006
Figure 28 (A) SST anomalies for the time period 1982–2006 (from http://iri.columbia.edu/climate/ENSO);
(B) average number of scallop Argopecten purpuratus settlers per collector bag (individuals bag−1) and the
number of collector bags placed in Bahia Tongoy (30°S); (C) harvest of Gracilaria chilensis between 18°S
and 35°S from natural banks and from culture beds. (Source for A and B: SERNAPESCA, Region IV, Chile.)
on the populations of large kelps and other macroalgae in northern Chile (Vásquez 1999). Kelp
aquaculture is presently in a developmental phase and has not yet achieved economic importance.
Other resources currently being investigated for aquaculture in northern Chile are bivalves from
sandy beaches (Mesodesma donacium, Tagelus dombeii). Culture of these bivalves is aiming at
stock repopulations after extinction of local stocks (see also Artisanal benthic fisheries, p. 278ff.).
Additionally, there are several introduced species that are presently cultured in northern Chile.
The Pacific oyster (Crassostrea gigas) was originally introduced in northern Chile in the 1970s,
and despite fast growth rates, aquaculture activities then moved to southern Chile because there
culture costs are cheaper (bottom culture vs. suspended culture in northern Chile). The production
in northern Chile is marginal, and there are only two oyster companies remaining. During the first
half of 2004 these produced only 935 t, which represents a 46.8% decrease compared with the
production during the first half of 2003. The influence of ENSO on the production of oysters is
not well known, but since Pacific oysters have a wide range of temperature tolerance EN effects
may be minor (or positive).
294
Other species introduced to the coast of Chile are abalone, originating from California and
Japan. These have been mainly cultured in land-based facilities, but due to an increasing production
and limited holding capacities on land, sea-based culture (as already established in southern Chile)
is also considered for northern-central Chile. The production of abalones increased from 1 t in
1998 to 342 t in 2005, and for the year 2006 it is expected that the Chilean abalone industry will
produce >500 t. Only in Region IV, currently five abalone production centres are established and
an additional five centres have solicited permits to initiate new aquaculture activities during 2006.
The present abalone production in Chile is mainly based on the red abalone (Haliotis rufescens).
However, the Japanese abalone (H. discus hannai), which has a better market value, is also raised,
but to a lesser degree since culture technology has higher requirements (and costs) than those for
red abalone.
Although the land-based abalone culture is not directly affected by variations in environmental
conditions, ENs may have severe effects on abalone culture because they can produce strong impacts
on the population of large kelp, the main food resource presently used in abalone culture. The lack
in supply of fresh food algae may produce serious bottlenecks in the culture of abalone. Some of
these problems are occurring presently and the National Fisheries Service (SERNAPESCA) is
concerned with the overexploitation of kelp, restricting the extraction from natural kelp beds and
promoting research for cultivation and management of seaweeds. This scenario presents important
challenges for applied research in the near future.
In general, aquaculture in northern and central Chile does not reach the levels it has in southern
Chile. Some of the main reasons for this are related to the fact that the coast of northern Chile is
(1) mostly exposed to wave action and (2) is strongly affected by important interannual variations
in oceanographic conditions. Future efforts should probably focus on the development of land-
based culture facilities and integrated systems where animals and algae are produced in combination
(Chopin et al. 2001).
Conservation of marine biodiversity

and Marine Protected Areas
The HCS extending from Ecuador to southern Chile is considered as one of the large marine
ecosystems for high-priority attention (Boersma et al. 2004). The growing use of coastal areas by
human activities is also increasingly threatening marine biodiversity in the HCS. The most important
threats are overfishing, aquaculture, pollution by sewage and mining activities, runoffs of chemicals
used for agriculture, oil spills and tourism activities (Vásquez et al. 1999, Fernández et al. 2000).
All countries, including Chile, that have ratified the Convention on Biological Diversity (CBD)
treaty agreed to develop a network of Marine Protected Areas (MPAs, as defined by IUCN 1994)
to ultimately protect 10% of the marine environments by 2012, based on an ecosystem approach
(Wood 2006). However, the actual establishment rate of MPAs (4.5% annual increase) reveals that
the work plan is unrealistic and will not be achieved before the second half of this century (Wood
2006). The Chilean case is a paradox as its economic exclusive zone (EEZ) corresponds to 17.8%
of the Latin-American EEZ and is three times larger than its terrestrial territory (~18% of which is
already protected; Pauchard & Villarroel 2002). However, only 0.03% of the Chilean EEZ (0.67%
of the territorial sea) is protected as MPAs (CONAMA personal communication). Further, only
14% of the surface area of these MPAs are located within the HCS of northern and central Chile
(18°S to 41°S). Considering that ~95% of the Chilean population is located between 18°S and 41°S
and the growing human impact in coastal areas, this zone represents one of the greatest challenges
for marine conservation.
The Chilean fisheries management policy, through creation of Management and Exploitation
Areas (AMERBs) for benthic resources from coastal habitats, is focusing on economically important
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MARTIN THIEL ET AL.
target species, largely ignoring habitats, ecological interactions and other important ecosystem
components (e.g., species dispersal). Although management areas can provide nursery grounds for
target species (e.g., the Chilean abalone, Concholepas concholepas, keyhole limpets Fissurella spp.
and the red sea urchin Loxechinus albus), this approach is often criticised for its weakness in
providing a long-term ecological and economic viability and uncertainty in its efficiency (NRC
2001). The establishment of MPAs has proven to be a useful tool to achieve conservation and
preservation goals (e.g., resources, communities, habitats), either as no-take MPAs or multiple-use
MPAs (MUMPAs) (NRC 2001). Among the different tools existing in the Chilean legislation for
protecting coastal areas, two types of no-take MPAs (marine reserves and parks) were foreseen in
the law since 1989 but have only recently been established (Morales & Ponce 1997, Fernández &
Castilla 2005). According to the Chilean laws, marine reserves are not focused on ecosystem
protection but rather on exploited resources and their habitats and eventually may allow partial
extractions if stocks reach very high levels of abundance. Three marine reserves are located between
18°S and 41°S (Table 6): La Rinconada to preserve a genetic stock of the scallop Argopecten
purpuratus and Isla Choros-Damas and Isla Chañaral for allowing the recovery of several over-
exploited benthic invertebrates (and their habitats). Furthermore, two MUMPAs, Isla Grande de
Atacama and Lafken Mapu Lahual, have been created by the National Environmental Agency
(CONAMA) with international private funding (GEF) as two of the three Chilean MUMPAs. In
contrast to the marine reserves, these MUMPAs aim at the conservation of biodiversity integrating
socioeconomic interests by creating not only no-take areas, but also areas where fishery and outdoor
activities (e.g., diving, ecotourism) are permitted. While no-take zones have not yet been established
in the MUMPA Isla Grande de Atacama, they have recently been identified (C. Gaymer et al.
unpublished data) and will be declared in the near future. Finally, a small no-take MPA is located
at Las Cruces, mainly for scientific purposes. Although very few MPAs exist in Chile, initiatives
for conservation are facilitated by the Chilean law, which established an exclusive zone for artisanal
fisheries within 5 nautical miles from the shore (e.g., prohibits trawling).
The MPAs Isla Choros-Damas, Isla Chañaral and Isla Grande de Atacama in northern Chile
have been subjectively chosen through the use of expert criteria, based on the presence of some
important ecological communities representing the HCS. The subtidal zone of these three MPAs
is characterised by kelp beds of Lessonia trabeculata, L. nigrescens and Macrocystis integrifolia
and several invertebrate populations overexploited during decades, and now vulnerable, such as the
Chilean abalone Concholepas concholepas and the red sea urchin Loxechinus albus (i.e., extraction
prohibited). Moreover, these MPAs are the habitat of some emblematic and/or endangered species,
such as the bottlenose dolphin Tursiops truncatus, the sea otter Lontra felina, the Humboldt penguin
Spheniscus humboldti and the Peruvian diving petrel Pelecanoides garnotii, and Isla Grande de
Atacama is the southernmost site where the wedge-rumped storm-petrel Oceanodroma tethys
kelsalli occurs. Biological corridors (Kaufman et al. 2004) permitting those species along with
other seabirds to travel between these three MPAs are not included in the present MPA design.
Additional MPAs between the marine reserves and the MUMPA would help to ensure undisturbed
migration of marine birds and mammals. Ultimately, given the use of both terrestrial and marine
environments by some species (e.g., Humboldt penguins) (Fariña et al. 2003b, Ellis et al. 2006),
the selection of priority sites for conservation/preservation should integrate both environments,
looking for common hot spots that would increase efficiency and reduce conservation costs.
MPAs offer a management tool to preserve hot spots of native species diversity; however, these
hot spots can be strongly affected by invasion of exotic species which could compromise the
effectiveness of MPAs (Byers 2005, Klinger et al. 2006). Introduction of invasive species in Chile
becomes a serious concern with the increase of aquaculture (Castilla et al. 2005a). The Isla Grande
de Atacama MPA is south of a bay (Bahia Inglesa) where intense scallop culture takes place and
where the highest density of the exotic seaweed Codium fragile for the Chilean coast has been
296
Table 6 Main characteristics of marine protected areas (MPAs) in the Humboldt Current System (HCS) between 18° and 41°S
Distance to Main
Establishment Mean Size next MPA conservation Main communities Biological
Name Status year latitude (km2) (km) to south goals Major threats and target groups survey
La Rinconada Marine 1997 23°28′ 3.4 404 Genetic reserve Illegal extractions Scallop bed Every year
Reserve S since 1997
Isla Grande de MUMPA 2004 27°10′ 35.5 213 Biodiversity Invasive species Kelp beds, barrens, 1 in 2002
Atacama S protection marine mammals
and birds
Isla Chañaral Marine 2005 29°02′ 4.3 20 Overexploited Illegal extractions Kelp beds, barrens, 1 in 1999
297
Reserve S species recovery marine mammals
and birds
Isla Choros-Damas Marine 2005 29°15′ 25 470 Overexploited Illegal extractions Kelp beds, barrens, 1 in 1999
Reserve S species recovery sea grass, marine
mammals and birds
Las Cruces No-take 2005 33°30′ 0.15 791 Scientific Illegal extractions Kelp beds, barrens Every year
MPA S research since 1982
Lafken Mapu Lahual MUMPA 2005 40°43′ 44.6 Biodiversity Conflicts with Kelp beds, marine 1 in 2006
S protection indigenous people mammals and birds
MARTIN THIEL ET AL.
reported (Neill et al. 2006). Recently, patches of C. fragile have been observed within the MPA
(R. Villablanca personal observations), highlighting the importance of considering connectivity
with surrounding areas and the constraints of aquaculture sites for selecting location of MPAs
(Micheli et al. 2004).
The establishment of MPAs in Chile has so far been based on anecdotal recommendations (e.g.,
resource management, tourist attractions) rather than scientific criteria. This approach is considered
inadequate to effectively protect biodiversity (Meir et al. 2004, Sutherland et al. 2004). Current
strategies for implementing MPA networks require a systematic planning conservation method to
identify optimal sites for protection of biodiversity (Sayre et al. 2000, Beck & Odaya 2001). The
first step in planning an MPA is the assessment and mapping (Geographic Information System —
GIS) of the coastal marine biodiversity, the physical environment and major threats (e.g., human
uses) and then the identification of priority sites using Decision Support Systems (DSS) based on
algorithms (Sala et al. 2002, Leslie et al. 2003). A DSS based on species richness has been used
to identify priority areas for marine vertebrate conservation along the Chilean coast (Tognelli et al.
2005). Habitat classification is generally considered as the conservation goal in the DSS (Roberts
et al. 2003). However, benthic surveys along the Chilean coast (C. Gaymer & C. Dumont unpub-
lished data) revealed that communities are probably more appropriate to characterise the benthic
environment and consider the ecosystem processes (e.g., trophic cascades; Shears & Babcock 2003),
ecological interactions (e.g., predator-prey; Micheli et al. 2004) and population connectivity (e.g.,
larval dispersal; Palumbi 2003). Moreover, there is an urgent need for more scientific information
in Chilean marine biodiversity (e.g., there is a lack of taxonomic expertise), population connectivity
(e.g., identifying source and sink populations) and ecological processes (in particular species
interactions in subtidal habitats are poorly studied).
Although ecological knowledge is a key component in developing MPAs, the management
effectiveness is the most important challenge for the success of an MPA (Mascia 2004, Pomeroy
et al. 2004). A major difficulty arises from the way in which marine reserves and MUMPAs have
been established in Chile. The former were created by an imposition from the central authority
(fisheries ministry) without consulting the stakeholders, who are mostly in disagreement with this
new status. This establishment strategy has turned enforcement into a complicated task for the
fisheries authority, and this may turn into a major threat for the success of present and future MPAs.
For example, since its creation in 1997, the marine reserve La Rinconada has been affected by
frequent illegal extractions of scallops (M. Avendaño personal observations). Social conflicts due to
lack of communication between the authority and the stakeholders are also present within the recently
created marine reserves Isla Choros-Damas and Isla Chañaral. In contrast, a participative process
took place in the establishment of the MUMPA Isla Grande de Atacama, incorporating most of the
relevant actors (i.e., administrative authorities, stakeholders, managers, scientists and fishermen),
offering the opportunity to evaluate contrasting interests in order to reduce potential conflicts. Social
costs should be evaluated before the establishment of MPAs and a formal educational process should
be implemented by the authorities to teach the importance of MPAs in developing sustainable
exploitation of resources (Mascia 2004). The government should also negotiate compensations and
propose alternative activities (e.g., tourism) to fishermen, who are the ancestral users of the MPA
areas, and avoid creating high expectations (Mascia 2004, Sobel & Dahlgren 2004).
Ideally, an international MPA network (from Ecuador to Chile) including the connectivity among
MPAs should be implemented to effectively preserve biodiversity in the HCS. This should be achieved
using the support of international tools and agreements, and international non-governmental organ-
isations (NGOs) in order to co-ordinate and improve the quality of scientific information and reduce
the costs (Balmford et al. 2004). Moreover, the Chilean government must contribute to funding for
implementation and functioning of MPAs as successful conservation experiences from all over the
298
world have demonstrated that self-funding (e.g., through tourism business) is not feasible (Balmford
et al. 2004).
Outlook, long-term research vision and future research frontiers

An outlook of the pressing scientific questions and tasks to be addressed in the short/mid-term
future (during the next decade) within the HCS of northern and central Chile should, as a minimum,
include (1) studies of ocean–atmosphere interactions and offshore oceanography, (2) research in
inshore and offshore oxygen-minimum ecosystems, (3) research on inner inshore coastal oceanog-
raphy and benthic-pelagic linkages, (4) development of an ecosystem-based adaptive resource
management approach to fisheries that integrates socioeconomic aspects, (5) implementation of a
coastal overarching network system for marine conservation-management, (6) novel approaches in
coastal mariculture, (7) studies of marine non-indigenous species (NIS), (8) marine molecular
biology, particularly on genomics, and (9) training of Chilean marine taxonomists.
A long-term HCS vision (during the coming two decades) should also, as a minimum, include
(1) intensification of precautionary and integrative ecosystem management; (2) implementation of
a high-sea conservation policy; (3) intensification of research on the continental slope, deep-sea
and abyssal ecosystems; (4) scientific and technological research on deep-sea gas (methane)
hydrates; and (5) evaluations of the effects of future climate change.
Short/mid-term scientific outlook

Research on ocean–atmosphere interactions and offshore oceanography
The HCS offers unique opportunities for offshore oceanographic studies (e.g., Strub et al. 1998)
and considered as a whole, Chile and Peru represent between 15% and 20% of the world’s fishery
landings. Large-scale fluctuations in ocean climate (ENSO and the PDO) dominate interannual and
interdecadal variability in the ocean, which in turn are linked to upwelling and climate changes
(Chavez et al. 2003) and are considered key elements in the HCS functioning. Upwelling systems
presently are experiencing ‘anomalous changes’ such as profound changes in the physical and
biogeochemical properties in the California Current Systems (Freeland et al. 2003, Grantham et al.
2004), massive nitrogen loss in the Benguela upwelling system (Kuypers et al. 2005), and hydrogen
sulphide eruptions in the Atlantic Ocean off southern Africa and linked abrupt degradation of
upwelling systems (Bakun & Weeks 2004, Weeks et al. 2004, Arntz et al. 2006). Such interannual
and decadal variability and anomalous changes may intensify due to global climate changes, which
will also affect the HCS, causing important changes in productivity, biogeochemical cycling and
fisheries. Research on these and related oceanographic topics is urgently needed for the HCS.
Research on inshore and offshore oxygen-minimum ecosystems
Oxygen-minimum zones (OMZs) in the ocean generally form along the EBCs. The decomposition
of upwelling-derived biomass in combination with sluggish circulation of mid-water masses
strongly enhance the hypoxia conditions, as is the case in the HCS (Levin & Gage 1998, Morales
et al. 1999, Levin 2002, Helly & Levin 2004, Ulloa & De Pol 2004). The HCS is characterised by
the relative shallowness of the oxygen-minimum layer. The OMZ produces peculiar environments
with organisms highly resistant to low oxygen concentrations (Levin et al. 2001, Gallardo et al.
2004). These environments are unique in the HCS and quite different from those off California
(Arntz et al. 2006). They offer diverse opportunities to develop frontier research, ranging from
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evolutionary adaptability, primary and secondary production, biodiversity, and species invasions
(Castilla & Neill 2007) to impacts on fisheries.
Research on nearshore coastal oceanography and benthic-pelagic coupling
The nearshore oceanography (coastal border to about 2–3 km offshore) is one of the least-known
areas of the world’s oceans. In this regard, during the past 5 yr several lines of research have been
developed in central Chile (e.g., Poulin et al. 2002a,b, Kaplan et al. 2003, Wieters et al. 2003,
Narváez et al. 2004, Vargas et al. 2004, Piñones et al. 2005) and further research efforts are needed.
Unless improvements in the knowledge of nearshore oceanography are achieved, coastal ecology
cannot be properly understood. This includes topics such as dispersal of nearshore propagules,
benthic resource fisheries and sustainability issues, coastal conservation and pollution impacts.
Linking nearshore oceanography and the coastal benthic-pelagic systems remains as a challenge,
not only along the HCS (Castilla et al. 2002a, Escribano et al. 2002, Lagos et al. 2002, Marín &
Moreno 2002), but also around the world (Shanks 1983, 1995, 2002, Largier 2002).
Development of an integrative and adaptive resource management

approach to fisheries
Pelagic, benthic and demersal marine resource management in Chile is regulated by the Ley de
Pesca y Acuicultura 1991 (Fishery and Aquaculture Law, FAL 1991), under the responsibility of the
Subsecretary of Fisheries, Ministry of Economy. For the management and administration of fish-
eries, among other regulations, the law (1) defines the ‘artisanal fishery’ referring to vessels/boats
<50 t and <18 m long, separating fishers into four major categories and distinguishing them from
the ‘industrial fisheries’; (2) establishes the allocation of a 5-mile wide coastal stretch from 18°20′S
to 41°28′S exclusively for the operation of the artisanal fishery fleet; (3) includes an artisanal and
industrial National Register Fisher System restricting the spatial movements of fleets; (4) promotes
co-management and allocates exclusive AMERBs and TURFs (for artisanal fishery associations);
and (5) regulates overexploited stocks via area closure systems and allocation of transferable and
non-transferable quotas (Defeo & Castilla 2005, Castilla & Gelcich 2006, Castilla et al. 2007).
Hence, it is herein suggested that the 1991 Chilean FAL contains modern and advanced fishery
regulation concepts and management tools to potentially promote the rationalisation of Chilean
fisheries. An important step forward in this direction must be the introduction of an integrated
socio-biophysical ecosystem fishery approach (FAO 2003, Castilla & Defeo 2005). To promote this
approach at least three elements need to be considered: (1) the proactive participation of stakeholders
(artisanal and industrial) in the co-evaluation of stocks, (2) a proactive government approach to
adaptive co-management (e.g., artisanal fisheries, AMERBs and TURFs) via the use of experimental
fisheries and further socioeconomic studies, and (3) an increase in the number of Chilean scientists
engaged in fishery biology and modelling of resource dynamics. In particular, this last point is
considered critical in achieving the stakeholder participation in stock evaluation and management
because well-trained scientists and communicators are needed to make information available trans-
parently, while simultaneously consulting and incorporating the perceptions of fishermen. Fostering
fisheries sciences would probably also enhance international collaboration, thereby also promoting
refereed frontier publications (which presently are extremely scant; Castilla et al. 2005b).
Implementation of a coastal overarching network system for marine

conservation management
A novel and overarching integrative approach to jointly address coastal conservation and manage-
ment issues along the Chilean coast needs to be implemented in the next decade. The aim will be
300
to build a coastal conservation management network, including no-take and multiple-use MPAs,
sanctuaries, marine concessions and AMERBs (Castilla 2000, Secretariat of the Convention on
Biological Diversity SCBD 2004, Fernández & Castilla 2005, Castilla & Gelcich 2006). The present
authors believe that, based on present progress, this overarching framework can be achieved in
Chile (following guidelines given by the SCBD 2004). In fact, along ~18–41°S at present there
are already established >300 AMERBs, two main MPAs for multiple uses (Isla Grande de Atacama
and Lafken Mapu Lahual; see CONAMA-PNUD 2006), four declared and active marine reserves
and several other categories of protected areas (Fernández & Castilla 2005). Connecting these areas
and providing them with one unified administrative umbrella will furthermore meet the goals of the
Chilean National Biodiversity Strategy Plan, which attempts to protect 10% of relevant marine
Chilean ecosystems by 2012 (Rovira 2006). This overarching network system must also consider
external threats such as pollution, which is an important issue in Chile due to pollution resulting
from mining activities, agriculture and ship paints (Correa et al. 1999).
Novel approaches in coastal mariculture
The Chilean coastline along the HCS is very exposed to the open ocean, generally lacking wave-
protected embayment, and is thus not particularly suitable for the development of mariculture
activities. Exceptions are the bays or bay systems of Mejillones, Antofagasta, Caldera, Coquimbo
and Dichato, where scallops, oysters, mussels and Gracilaria are cultured on a limited scale. Inland
mariculture has also been developed, particularly for introduced species of the genus Haliotis. If
further progress for mariculture in the HCS will occur, the challenge remains in the development
of novel technologies for sea bottom or raft culture systems in exposed and offshore systems.
Research on marine non-indigenous species
The number of NIS along the HCS is surprisingly low when compared with similar upwelling or
non-upwelling systems around the world (Castilla et al. 2005a). It has been suggested that this
results from a combination of factors such as less-stressed coastal environments or the scarcity of
estuaries, gulfs and enclosed bays. Furthermore, it has been hypothesised that it might be linked
to the existence of the coastal shallow oxygen-minimum zones (Castilla & Neill 2007). These
aspects need further research. Moreover, the rate at which NIS are presently being introduced for
aquaculture purposes (e.g., salmon, abalone, algae) needs to be carefully monitored since, for
instance, escapees (e.g., salmon) from culture pens may impact native species and communities
(Buschmann et al. 2006b). The potential for the introduction of diseases and pests into HCS coastal
environments, linked to the development of aquaculture (e.g., Radashevsky & Olivares 2005, R.A.
Moreno et al. 2006b, Neill et al. 2006), must also be monitored.
Research in marine molecular biology, particularly on genomics
The use of molecular biology techniques in marine organisms is seen for Chile as a new research
frontier (Castilla et al. 2005b), allowing a range of questions to be addressed, covering evolutionary
biology, taxonomy and phylogeny (e.g., Letelier et al. 2003, Véliz et al. 2003, Thiel et al. 2004),
ecology and invasive species (Castilla et al. 2002b) and above all population aspects linked to
fisheries (Cárdenas et al. 2005). We anticipate that during the next decade this research, particularly
on marine genomics, should substantially increase.
Training of Chilean marine taxonomists
In Chile the training of marine taxonomists must increase. For this the establishment of specific
programmes is needed that support the formation and hiring of taxonomic experts, publication of
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monographs and field guides, visiting taxonomist programmes with the objective to tackle certain
groups and provide publications that allow easy identification of the HCS species, visiting scholar-
ships for national scientists to foreign institutions and museums, increased support for functioning
species collections, and an innovative programme to link specimen collections and molecular infor-
mation. The enhancement of marine taxonomic research in Chile embodies the vision that biodiver-
sity and ecological functioning are key components of the HCS. If this ecosystem is going to be
fully understood, and fishery, environmental sustainability and conservation programmes are to be
implemented, then the improvement of taxonomic expertise is seen as crucial.
HCS long-term scientific vision

Intensification of precautionary and integrative ecosystem management
A consideration of the research needs for the HCS indicates that fishery sustainability (subsistence,
small-scale and industrial fisheries) will continue for a long time to play a critical role in the
economy and human well-being of Chile. Presently, the FAO Code of Conduct Fishery Precaution-
ary Principles and the socio-biophysical ecosystems fishery management approach (FAO 1995,
2003) are the two most novel management tools regarding sustainability of world fisheries. Fol-
lowing the failure of traditional single-species approaches (Defeo et al. 2007), the present authors
consider these tools as key elements for an efficient fisheries management in the HCS, and scientists
in Chile need to be trained and prepared to properly use them. Therefore, an intensification of
scientific research in these areas with respect to the HCS is foreseen. There is a particular need for
communication between scientists, users, administrators and politicians. Due to the overriding
importance of the HCS for the Chilean society (fisheries, aquaculture, shipping, tourism), integrative
management requires one unifying administrative body that oversees, monitors and evaluates all
activities within the HCS of Chile. This entity should incorporate representatives of all interest
groups and co-ordinate communication among them on both the national and international levels.
Implementation of a high-sea conservation policy
So far, Chile has not developed a high-sea conservation policy, and there is no policy for unique
offshore oceanic realms, such as sea mountains and deep-sea environments, or populations of
migratory mammals, birds and fishes. Extensive and international protection measures for highly
mobile, migratory marine vertebrates in the dynamic high sea of the HCS are needed. The imple-
mentation of such policies remains an important challenge for Chile.
Intensification of the research on continental slope, deep-sea and abyssal ecosystems
The Chilean portion of the continental slope, deep-sea and abyssal realm of the HCS continue to
be some of the least-known oceanographic ecosystems. Aside from isolated studies (e.g., Glud et al.
1999, Thurston et al. 2002), little is known about the OM accumulation, benthic ecology or species
diversity in the unique deep trenches off the Chilean coast. During the next two decades or so Chile
has to make an effort to improve the understanding of the HCS trenches and also to increase
research in offshore oceanography. In order to achieve this goal, the country urgently needs to
improve access to oceanic work platforms, including modern research vessels (Castilla et al. 2005b).
Scientific and technological research on deep-sea gas (methane) hydrates
Gas hydrates are solid crystals formed by a cell of water molecules containing methane, ethane,
CO2 or H2S and are found inside pores of sedimentary rocks. Methane hydrates are considered to
302
be one of the strategic energy compounds for the future (they commonly occur below the permafrost
shield and in sediments of ocean margins). Gas methane hydrates are abundant on the continental
margins between Papudo (central Chile) to Valdivia (southern Chile). First evaluations in this area
indicate gas methane hydrate reserves for 1013 m3 (Morales 2003). The technology to access these
hydrates is developing fast, and Chile must be part of the advance and innovation processes.
Geological oceanography in Chile is poorly developed (although the number of geologists in the
country is high). Therefore the training of marine geologists and the substantial increase of marine
geological research in Chile remains an important task (and opportunity) for the future (Castilla
et al. 2005b).
Evaluation of the effects of future climate change
It is known that greenhouse warming and other human alterations may increase the possibility of
large and abrupt regional or global changes in climatic events, oceanic circulation (especially related
to deep-water formation), sea–ice dynamics (Smith et al. 2006), and wind velocity (Bakun & Weeks
2004). Future wind increases, due to greenhouse effects, may eventually affect one of the HCS key
characteristics, such as the rate of upwelling events (Bakun 1990, Bakun & Weeks 2004). For
instance, an increase of 15% in wind would represent an increase of ~40% in the typical rate (late
twentieth century) of sea upper layer volume replaced per day (water renewal). In the Benguela
Current System off Namibia, the present atmospheric greenhouse-related intensification of coastal
upwelling appears to be causing the abrupt degradation of the ecosystem (Weeks et al. 2004).
Therefore, in a scenario with concentrations of greenhouse gases increasing, climate change (e.g.,
pattern of wind increase and intensification of upwelling) along the HCS needs to be surveyed
under a long-term monitoring scheme.
HCS frontier research opportunities

As pointed out there are several research opportunities along the HCS, such as studies of the OMZ
and related ecosystems, upwelling and climate change, fishery productivity, co-management, con-
servation and socio-biophysical approaches to fisheries. In addition there are other very exciting
and novel frontier lines of research such as studies of the biogeochemical role of Archaea (Woese
& Fox 1977) in the HCS. These organisms are also linked to the OMZ and appear to be particularly
abundant in this system. This line of research has recently started in Chile and needs to be
strengthened.
Further topics that extend far beyond the scope of the present review but that appear to be
relevant in the present context are linked to socioecology, anthropology and paleo-oceanography.
For example, how do the processes occurring in coastal waters of the HCS affect the terrestrial
environments and human populations along the Pacific coast of South America (past, present and
future)? Palaeo-oceanography and anthropology can offer exciting insights from the past that will
also help to master future challenges.
Conclusions
Upwelling is the major driving force of ecological processes in the HCS by promoting high PP
both in the plankton and in the nearshore benthos. Additional processes influenced by upwelling
are transport of propagules and biogeochemical processes. Besides their high nutrient concentra-
tions, one main feature of the upwelled waters from the HCS along the Chilean coast is that they
have low concentrations of dissolved oxygen. This restricts vertical migration of most zooplankters,
including of larvae, thereby affecting dispersal. Furthermore, the low oxygen concentrations drive
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the dynamics of the benthos communities along the continental shelf and also the remineralisation
of OM. There is indication that the upper depth limits of the low-oxygen fauna oscillates with
upwelling strength (and EN). In parallel with these oscillations, the lower depth limit of benthos
communities from the upper sublittoral zone shifts up and down. This community experiences at
its upper depth limit the impact of EN events, where individual species or the whole community
may temporarily disappear (due to water temperature, wave action or burial under terrigenous
sediments). These extinction events affect the shallow sublittoral and intertidal biota along the HCS
in northern and central Chile, but their intensity varies between events, often resulting in differential
elimination of particular species, while leaving others unaffected or favouring them. Similar vari-
ation in effect size is seen along the latitudinal gradient, where EN impacts (which may occur at
all trophic levels) attenuate toward higher latitudes. This leads to different constellations in the
interaction webs found in pelagic and benthic communities. Spatial variability in upwelling, which
mitigates EN effects, further enhances the variability in the ecological responses. The diffuse pattern
of biogeographic limits observed in northern-central Chile is expression of this high variability.
Studies of individual growth, reproductive potential, dispersal, recruitment, physiology and popu-
lation connectivity of organisms from the HCS demonstrate the importance of life-history traits for
improving predictability of ecological processes in this area. The high temporal and spatial vari-
ability in oceanographic conditions and ecological processes in the HCS of northern-central Chile
complicates management and conservation decisions (such as calculation of catch quotas or iden-
tification of important areas for recruitment or growth). Recent studies have revealed small-scale
variability in reproductive potential, larval supply, recruitment and growth. This indicates that
spatially explicit conservation measures (e.g., MPAs) require information of high temporal and
spatial resolution. For most parts of the coast of northern and central Chile such data are not
available. Given this present gap in information and the urgent need for efficient conservation of
this large marine ecosystem, a large-scale approach is proposed. Future conservation measures
should also include terrestrial environments such as seabird breeding sites, dune fields and estuarine
habitats. Research and administration activities along the HCS face important challenges that require
substantial efforts, in particular a continuous and fluent communication among all involved parties.
In order to achieve this, a common umbrella organisation that co-ordinates all these activities (and
permits rapid exchange of opinion and information) appears to be highly desirable.
Acknowledgements
We acknowledge the continuing financial support from FONDECYT, FONDAP, FONDEF, FIP,
SHOA, MECESUP and DAAD. We are especially grateful to Jim Atkinson for his patience and
careful editing of the manuscript. We also would like to thank all our colleagues who allowed us
to refer to their unpublished data and manuscripts.
References
Aburto, J. & Stotz, W. 2003. Una experiencia de co-manejo de bivalvos en el marco de una nueva herramienta
de administración pesquera en Chile: las áreas de manejo. Policy Matters 12, 200–204.
Acuña, E., Arancibia, A., Mujica, A., Cid, L. & Roa, R. 1998. Análisis de la pesquería y evaluación indirecta
del stock de langostino amarillo en la III y IV Regiones. Informe Final FIP No. 96-08. Coquimbo,
Chile.
Acuña, E., Arancibia, A., Roa, R., Alarcón, R., Díaz, C., Mujica, A., Winkler, F.M. & Lépez, I. 1997. Análisis
de la pesquería y evaluación indirecta del stock del camarón nailon (II a VIII Regiones). Informe
Final FIP No. 95-06. Coquimbo, Chile.
304
Acuña, E., Moraga, J. & Uribe, E. 1989. La zona de Coquimbo: un sistema nerítico de surgencia de alta
productividad. In Memorias del Simposio Internacional de los Recursos Vivos y las Pesquerías en el
Pacífico Sudeste, R. Jordán et al. (eds). Santiago, Chile: Comité Permanente Pacífico Sur, 145–157.
Acuña, E., Villarroel, J.C., Bodini, A. & Andrade, M.J. 2005. The yellow squat lobster (Cervimunida johni)
fishery off northern Chile, 1994–2001. In Deep Sea 2003: Conference on the Governance and
Management, R. Shotton (ed.). Rome: FAO Fisheries Proceeding No. 3/2, 79–84.
Aguilar, A., Reid, C. & Thorpe, A. 2000. The political economy of marine fisheries development in Perú,
Chile and México. Journal of Latin American Studies 32, 503–527.
Aguilar, M. & Stotz, W. 2000. Settlement sites of juvenile scallops Argopecten purpuratus (Lamarck, 1819)
in the subtidal zone at Puerto Aldea, Tongoy Bay, Chile. Journal of Shellfish Research 19, 749–755.
Ahumada, R. 1989. Producción y destino de la biomasa fitoplantónica en un sistema de bahías en Chile central:
una hipótesis. Biología Pesquera 18, 53–66.
Ahumada, R., Pinto, L. & Camus, P.A. 2000. The Chilean coast. In Seas at the Millenium: an Environmental
Analysis, C.R.C. Sheppard (ed.). Oxford, U.K.: Pergamon Press, 699–717.
Ahumada, R., Rudolph, A. & Martínez, V. 1983. Circulation and fertility of water in Concepción Bay, Chile.
Estuarine Coastal and Shelf Science 16, 95–105.
Aiken, C., Navarrete, S.A., Castillo, M. & Castilla, J.C. 2007. Along-shore larval dispersal kernels in a
numerical ocean model of the central Chilean coast. Marine Ecology Progress Series (in press).
Alarcón, E. & Navea, M. 1992. Estudio del recurso de machas (Mesodesma donacium) en la IV Región.
Informe Final. SERPLAC, La Serena, Chile.
Alheit, J. 1987. Egg cannibalism versus egg predation: their significance in anchovies. South African Journal
of Marine Science 5, 467–470.
Alheit, J. & Niquen, M. 2004. Regime shifts in the Humboldt Current ecosystem. Progress in Oceanography
60, 201–222.
Alvarado, J.L. & Castilla, J.C. 1996. Tridimensional matrices of mussels Perumytilus purpuratus on intertidal
platforms with varying wave forces in central Chile. Marine Ecology Progress Series 133, 135–141.
Alvarado, J.L., Pinto, R., Marquet, P., Pacheco, C., Guiñez, R. & Castilla, J.C. 2001. Patch recolonization by
the tunicate Pyura praeputialis in the rocky intertidal of the Bay of Antofagasta, Chile: evidence for
self-facilitation mechanisms. Marine Ecology Progress Series 224, 93–101.
Alveal, K., Romo, H. & Valenzuela, J. 1973. Consideraciones ecológicas de las regiones de Valparaíso y
Magallanes. Revista de Biología Marina, Valparaíso (Chile) 15, 1–29.
Angel, A. & Ojeda, F.P. 2001. Structure and trophic organization of subtidal fish assemblages on the northern
Chilean coast: the effect of habitat complexity. Marine Ecology Progress Series 217, 81–91.
Antezana, T. 1978. Distribution of euphausiids in the Chile-Perú Current with particular reference to the
endemic Euphausia mucronata and the oxygen minimum layer. Ph.D. dissertation, University of
California, San Diego.
Antezana, T. 1981. Zoogeography of euphausiids in the south eastern Pacific Ocean. In Memorias del Seminario
sobre Indicadores Biológicos del Plancton, M. Vega (ed.). Montevideo, Uruguay: UNESCO, 5–23.
Antezana, T. 2002. Vertical distribution and diel migration of Euphausia mucronata in the oxygen minimum
layer of the Humboldt Current. In Oceanography of the Eastern Pacific II, J. Farber (ed.). Ensenada,
Mexico: CICESE, 13–28.
Arana, P. & Nakanishi, A. 1971. La pesquería del camarón nailon (Heterocarpus reedi) frente a la costa de
Valparaíso. Investigaciones Marinas, Valparaíso 2, 61–92.
Arancibia, H., Cubillos, L.A. & Acuña, E. 2005. Annual growth and age composition of the squat lobster
Cervimunida johni off northern-central Chile (1996–1997). Scientia Marina 69, 113–122.
Arancibia, H. & Neira, S. 2005. Long-term changes in the mean trophic level of central Chile fishery landings.
Scientia Marina 69, 295–300.
Arata, J. & Xavier, J.C. 2003. The diet of black-browed albatrosses at the Diego Ramirez Islands, Chile. Polar
Biology 26, 638–647.
Arcos, D.F., Cubillos, L.A. & Núñez, S.P. 2001. The jack mackerel fishery and El Niño 1997–1998 effects
off Chile. Progress in Oceanography 49, 597–617.
Arntz, W. 1986. The two faces of El Niño 1982–1983. Meeresforschung 31, 1–46.
305
MARTIN THIEL ET AL.
Arntz, W.E., Brey, T., Tarazona, J. & Robles, A. 1987. Changes in the structure of a shallow sandy-beach
community in Peru during an El-Niño event. South African Journal of Marine Science 5, 645–658.
Arntz, W. & Fahrbach, E. 1991. El Niño — Klimaexperiment der Natur. Physikalische Ursachen und biolo-
gische Folgen. Basel: Birkhäuser Verlag.
Arntz, W.E., Gallardo, V.A., Gutiérrez, D., Isla, E., Levin, L.A., Mendo, J., Neira, C., Rowe, G.T., Tarazona,
J. & Wolff, M. 2006. El Niño and similar perturbation effects on the benthos of the Humboldt,
California, and Benguela Current upwelling ecosystems. Advances in Geosciences 6, 243–265.
Arntz, W., Tarazona, J., Gallardo, V.A., Flores, V.A. & Salzwedel, H. 1991. Benthos communities in oxygen
deficient shelf and upper slope areas of the Peruvian and Chilean Pacific coast, and changes caused
by El Niño. In Modern and Ancient Continental Shelf Anoxia, R.V. Tyson & T.H. Pearson (eds).
London: Geological Society Special Publications, Vol. 58, 131–154.
Arntz, W.E., Valdivia, E. & Zeballos, J. 1998. Impact of El Niño 1982–83 on the commercially exploited
invertebrates (mariscos) of the Peruvian shore. Meeresforschung 32, 3–22.
Asmus, R.M. & Asmus, H. 1991. Mussel beds limiting or promoting phytoplankton. Journal of Experimental
Astorga, A., Fernández, M., Boschi, E.E. & Lagos, N. 2003. Two oceans, two taxa and one mode of
development: latitudinal diversity patterns of South American crabs and test for possible causal
processes. Ecology Letters 6, 420–427.
Atkinson, L.P., Valle-Levinson, A., Figueroa, D., De Pol-Holz, R., Gallardo, V.A., Schneider, W., Blanco, J.L.
& Schmidt, M. 2002. Oceanographic observations in Chilean coastal waters between Valdivia and
Concepción. Journal of Geophysical Research-Oceans 107, C7, DOI 10.1029/2001JC000991.
Avaria, S. & Muñoz, P. 1982. Producción actual, biomasa y composición específica del fitoplancton de la
Bahía de Valparaíso en 1979. Revista de Biología Marina, Valparaíso (Chile) 18, 129–157.
Avaria, S., Muñoz, P. & Uribe, E. 1982. Composición y biomasa del fitoplancton marino del norte de Chile
en Diciembre de 1980 (Operación oceanográfica MARCHILE XI-ERFEN II). Ciencia y Tecología
del Mar CONA 6, 5–36.
Avendaño, M. & Cantillánez, M. 2005. Crecimiento y estructura demográfica de Argopecten purpuratus en
la reserva marina La Rinconada, Antofagasta, Chile. Ciencias Marinas 31, 491–503.
Avila, M., Santelices, B. & McLachlan, J. 1986. Photoperiod and temperature regulation of the life history
of Porphyra columbina (Rodophyta, Bangiales) from central Chile. Canadian Journal of Botany 64,
1867–1872.
Aviléz, O. & Jerez, G. 1999. Gestión sustentable de recursos marinos bentónicos en caletas de la IV Región.
Ambiente y Desarrollo (Chile) 15, 6–10.
Báez, P., Arata, J. & Jackson, D. 2004. El loco Concholepas concholepas (Bruguiere, 1789) (Mollusca:
Gastropoda: Muricidae) during the early-middle Holocene of Los Vilos, central Chile. Investigaciones
Marinas, Valparaíso 32, 107–113.
Báez, P., Quiroz, D. & Jackson, D. 1994. Crustáceos en la pesca prehispánica de Chile: antecedentes
arqueológicos. Chile Pesquero 81, 48–52.
Baeza, J.A. & Fernández, M. 2002. Active brood care in Cancer setosus (Crustacea: Decapoda): the relationship
between female behaviour, embryo oxygen consumption and the cost of brooding. Functional Ecology
16, 241–251.
Bakun, A. 1990. Global climate change and intensification of coastal ocean upwelling. Science 247, 198–201.
Bakun, A. 2001. ‘School-mix feedback’: a different way to think about low frequency variability in large
mobile fish populations. Progress in Oceanography 49, 485–511.
Bakun, A. & Nelson, C.S. 1991. The seasonal cycle of wind-stress curl in subtropical Eastern Boundary
Current regions. Journal of Physical Oceanography 21, 1815–1834.
Bakun, A. & Weeks, S.J. 2004. Greenhouse gas buildup, sardines, submarine eruptions and the possibility of
abrupt degradation of intense marine upwelling ecosystems. Ecology Letters 7, 1015–1023.
Balbontín, F., Llanos, A. & Valenzuela, V. 1979. Estudio experimental sobre selección de alimento y compor-
tamiento alimentario en anchoveta y sardina de Chile (Pisces, Clupeiformes). Revista de Biología
Marina, Valparaíso (Chile) 16, 211–220.
Balbontín, F., Llanos, A. & Valenzuela, V. 1997. Trophic overlap and feeding incidence in fish larvae from
central Chile. Revista Chilena de Historia Natural 70, 381–390.
306
Balmford, A., Gravestock, P., Hockley, N., McClean, C.J. & Roberts, C.M. 2004. The worldwide costs of
Marine Protected Areas. Proceedings of the National Academy of Sciences USA 101, 9694–9697.
Balech, E. 1954. División zoogeográfica del litoral sudamericano. Revista de Biología Marina, Valparaíso
(Chile) 4, 184–195.
Baums, I.B., Miller, M.W. & Hellberg, M.E. 2005. Regionally isolated populations of an imperiled Caribbean
coral, Acropora palmata. Molecular Ecology 14, 1377–1390.
Beck, M.W. & Odaya, M. 2001. Ecoregional planning in marine environments: identifying priority sites for
conservation in the northern Gulf of Mexico. Aquatic Conservation: Marine and Freshwater Ecosys-
tems 11, 235–242.
Becker, J. & Craig, E.A. 1994. Heat-shock proteins as molecular chaperones. European Journal of Biochem-
istry 219, 11–23.
Bernal, M., Simeone, A. & Flores, M. 2006. Breeding of wedge-rumped storm-petrels (Oceanodroma tethys)
in northern Chile. Ornitologia Neotropical 17, 283–287.
Bernal, P.A., Ahumada, R., González, H., Pantoja, S. & Troncoso, A. 1989. Flujo de energía en un modelo
trófico en la Bahía de Concepción. Biología Pesquera 18, 5–14.
Bernal, P.A., Oliva, D., Aliaga, B. & Morales, C. 1999. New regulations in Chilean fisheries and aquaculture:
ITQ’s and territorial users rights. Ocean and Coastal Management 42, 119–142.
Bertrand, A., Barbieri, M.A., Cordova, J., Hernández, C., Gómez, F. & Leiva, F. 2004. Diel vertical behaviour,
predator-prey relationships, and occupation of space by jack mackerel (Trachurus murphyi) off Chile.
ICES Journal of Marine Science 61, 1105–1112.
Bertrand, A., Barbieri, M.A., Gerlotto, F., Leiva, F. & Cordova, J. 2006. Determinism and plasticity of fish
schooling behaviour as exemplified by the South Pacific jack mackerel Trachurus murphyi. Marine
Bird, J. 1943. Excavations in northern Chile. Anthropological Papers of the American Museum of Natural
History 38, 173–318.
Bird, J. 1946. The cultural sequence of the northern Chilean coast. In Handbook of South American Indians,
J.H. Steward (ed.). Washington, D.C.: Smithsonian Institution, 587–594.
Bischof, K., Gómez, I., Molis, M., Hanelt, D., Karsten, U., Lüder, U., Roleda, M.Y., Zacher, K. & Wiencke,
C. 2006. Ultraviolet radiation shapes seaweed communities. Reviews in Environmental Science and
Biotechnology 5, 141–166.
Blanco, J.L., Thomas, A.C., Carr, M.E. & Strub, P.T. 2001. Seasonal climatology of hydrographic conditions
in the upwelling region off northern Chile. Journal of Geophysical Research-Oceans 106, 11,451–11,467.
Bobadilla, M. & Santelices, B. 2004. A new technique for simultaneous collection of macroalgal propagules
in the water column. Journal of Experimental Marine Biology and Ecology 298, 125–131.
Bobadilla, M. & Santelices, B. 2005. Variations in the dispersal curves of macroalgal propagules from a source.
Boersma, D., Ogden, J., Branch, G., Bustamante, R.H., Compagna, C., Harris, G. & Pikitch, E.K. 2004. Lines
on the water ocean-use planning in large marine ecosystems. In Defying Ocean’s End: and Agenda
for Action, L.K. Glover & S.A. Earle (eds). Washington, D.C.: Island Press, 125–138.
Boltovskoy, E. 1964. Províncias zoogeográficas de América del sur y su sector antártico según los foraminiferos
bentónicos. Boletín del Instituto de Biología Marina de Mar del Plata 7, 93–99.
Bomkamp, R.E., Page, H.M. & Dugan, J.E. 2004. Role of food subsidies and habitat structure in influencing
benthic communities of shell mounds at sites of existing and former offshore oil platforms. Marine
Biology 146, 201–211.
Botsford, L.W., Moloney, C.L., Hastings, A., Largier, J.L., Powell, T.M., Higgins, K. & Quinn, J.F. 1994. The
influence of spatially and temporally varying oceanographic conditions on meroplanktonic metapop-
ulations. Deep-Sea Research II 41, 107–145.
Böttjer, D. & Morales, C.E. 2005. Microzooplankton grazing in a coastal embayment off Concepción, Chile
(~36°S) during non-upwelling conditions. Journal of Plankton Research 27, 383–391.
Bowden, K.F. 1983. Physical Oceanography of Coastal Waters. Chichester, U.K.: Ellis Horwood.
Brandhorst, W. 1971. Condiciones oceanográficas estivales frente a la costa de Chile. Revista Biología Marina
14, 45–84.
307
MARTIN THIEL ET AL.
Brante, A., Cifuentes, S., Pörtner, H.O., Arntz, W. & Fernández, M. 2004. Latitudinal comparisons of reproductive
traits in five brachyuran species along the Chilean coast. Revista Chilena de Historia Natural 77, 15–27.
Brante, A., Fernández, M., Eckerle, L., Mark, F., Pörtner, H.O. & Arntz, W. 2003. Reproductive investment
in the crab Cancer setosus along a latitudinal cline: egg production, embryo losses and embryo
ventilation. Marine Ecology Progress Series 251, 221–232.
Brattström, H. & Dahl, E. 1951. General account, list of stations, hydrography. Reports of the Lund University
Chile Expedition 1948–1949. Lunds Universitets Arsskrift N.F. 46, 1–88.
Brattström, H. & Johanssen, A. 1983. Ecological and regional zoogeography of the marine benthic fauna of
Chile. Sarsia 68, 289–339.
Brey, T. 1995. Temperature and reproductive metabolism in macrobenthic populations. Marine Ecology
Briones, L., Núñez, L. & Standen, V.G. 2005. Geoglifos y tráfico prehispánico de caravanas de llamas en el
desierto de Atacama (norte de Chile). Chungará (Arica) 37, 195–223.
Broitman, B.R., Navarrete, S.A., Smith, F. & Gaines, S.D. 2001. Geographic variation of southeastern Pacific
intertidal communities. Marine Ecology Progress Series 224, 21–34.
Brokordt, K.B., Fernández, M. & Gaymer, C.F. 2006. Domestication reduces the capacity to escape from
predators. Journal of Experimental Marine Biology and Ecology 329, 11–19.
Brokordt, K.B., Himmelman, J.H. & Guderley, H.E. 2000a. Effect of reproduction on escape responses and
muscle metabolic capacities in the scallop Chlamys islandica Muller 1776. Journal of Experimental
Brokordt, K.B., Himmelman, J.H., Nusetti, O.A. & Guderley, H.E. 2000b. Reproductive investment reduces
recuperation from exhaustive escape activity in the tropical scallop Euvola zizac. Marine Biology 137,
857–865.
Brown, J.H. & Lomolino, M.V. 1998. Biogeography. Sunderland, Massachusetts: Sinauer Associates.
Bulboa, C.R. & Macchiavello, J.E. 2001. The effects of light and temperature on different phases of the life
cycle in the carrageenan producing alga Chondracanthus chamissoi (Rhodophyta, Gigartinales).
Botanica Marina 44, 371–374.
Burton, R.S. 1998. Intraspecific phylogeography across the point Conception biogeographic boundary. Evo-
lution 52, 734–745.
Buschmann, A.H. 1990. Intertidal macroalgae as refuge and food for amphipoda in central Chile. Aquatic
Botany 36, 237–245.
Buschmann, A.H., García, C., Espinoza, R., Filún, L. & Vásquez, J.A. 2004a. Sea urchin (Loxechinus albus)
and kelp (Macrocystis pyrifera) in protected areas in southern Chile. In Sea Urchins: Fisheries and
Ecology, J.M. Lawrence & O. Guzmán (eds). Lancaster, Pennsylvania: DEStech Publications, 120–130.
Buschmann, A.H., Moreno, C., Vásquez, J.A. & Hernández-González, M.C. 2006a. Population and reproduc-
tion strategies of Macrocystis pyrifera (Phaeophyta) in southern Chile. Journal of Applied Phycology
18, 575–582.
Buschmann, A.H., Riquelme, V.A., Hernández-González, M.C., Varela, D., Jiménez, J.E., Henríquez, L.A.,
Vergara, P.A., Guíñez, R. & Filún, L. 2006b. A review of the impacts of salmonid farming on marine
coastal ecosystems in the southeast Pacific. ICES Journal of Marine Science 63, 1338–1345.
Buschmann, A.H., Vásquez, J.A., Osorio, P., Reyes, E., Filun, L., Hernández-González, M.C. & Vega, A.
2004b. The effect of water movement, temperature and salinity on abundance and reproductive patterns
of Macrocystis spp. (Phaeophyta) at different latitudes in Chile. Marine Biology 145, 849–862.
Bustamante, R.H. & Branch, G.M. 1996. The dependence of intertidal consumers on kelp-derived organic
matter on the west coast of South Africa. Journal of Experimental Marine Biology and Ecology 196,
1–28.
Bustamante, R.H., Branch, G.M. & Eekhout, S. 1995. Maintenance of an exceptional intertidal grazer biomass
in South Africa — subsidy by subtidal kelps. Ecology 76, 2314–2329.
Butman, C.A. 1987. Larval settlement of soft-sediment invertebrates: the spatial scales of pattern explained
by habitat selection and the emerging role of hydrodynamical processes. Oceanography and Marine
Biology: An Annual Review 25, 113–165.
Byers, J.E. 2005. Marine reserves enhance abundance but not competitive impacts of a harvested nonindigenous
species. Ecology 86, 487–500.
308
Cáceres, H. 2001. Resolución taxonómica necesaria para evaluar los efectos de la contaminación provocados
por el vertimiento de aguas servidas sobre la fauna asociada a Gelidium chilense (Montagne)
Santelices y Montalva 1983 en el sector de La Pampilla Coquimbo. Honors thesis, Universidad
Católica del Norte, Coquimbo, Chile.
Cáceres, M. 1992. Vórtices y filamentos observados en imágenes satelitales frente al área de surgencias de
Talcahuano, Chile central. Investigaciones Pesqueras (Chile) 37, 55–66.
Cáceres, M. & Arcos, D. 1991. Variabilidad en la estructura especio-temporal de un área de surgencia frente
a la costa de Concepción, Chile. Investigaciones Pesqueras (Chile) 36, 27–38.
Caillaux, L.M. & Stotz, W.B. 2003. Distribution and abundance of Rhynchocinetes typus (Crustacea: Deca-
poda), in different benthic community structures in northern Chile. Journal of the Marine Biological
Camus, P.A. 1990. Procesos regionales y fitogeografía en el Pacífico Suroriental: el efecto de ‘El Niño-
Oscilación del Sur’. Revista Chilena de Historia Natural 63, 11–17.
Camus, P.A. 1994a. Recruitment of the intertidal kelp Lessonia nigrescens Bory in northern Chile: successional
constraints and opportunities. Journal of Experimental Marine Biology and Ecology 184, 171–181.
Camus, P.A. 1994b. Dinámica geográfica en poblaciones de Lessonia nigrescens Bory (Phaeophyta) en el
norte de Chile: Importancia de la extinción local durante eventos El Niño de gran intensidad. Revista
de Investigaciones Científicas y Tecnológicas, Serie Ciencias Marinas 3, 58–70.
Camus, P.A. 2001. Marine biogeography of continental Chile. Revista Chilena de Historia Natural 74, 587–617.
Camus, P.A. & Andrade, Y.N. 1999. Diversity of rocky intertidal communities in northern Chile and the
potential effect of coastal upwelling. Revista Chilena de Historia Natural 72, 389–410.
Camus, P.A. & Lagos, N.A. 1996. Variación espacio-temporal del reclutamiento en ensambles intermareales
sésiles del norte de Chile. Revista Chilena de Historia Natural 69, 193–204.
Camus, P.A., Vásquez, J.A., González, E.O. & Galáz, L.E. 1994. Fenología espacial de la diversidad comu-
nitaria intermareal en el norte de Chile: patrones comunitarios de variación geográfica e impacto de
los procesos de extinción-recolonización post El Niño 82/83. Medio Ambiente (Chile) 12, 129–163.
Cane, M.A. 2005. The evolution of El Niño, past and future. Earth and Planetary Science Letters 230, 227–240.
Capella, J., Vilina, Y. & Gibbons, J. 1999. Observación de cetáceos en isla Chañaral y nuevos registros para
el área de la reserva nacional Pingüino de Humboldt, Norte de Chile. Estudios Oceanologicos 18,
57–64.
Carcelles, A. & Williamson, S. 1951. Catálogo de los moluscos marinos de la provincia Magellánica. Revista
del Instituto Nacional de Investigaciones y Ciencias Naturales Bernardino Rivadavia 2, 225–283.
Cárdenas, L., Hernández, C.E., Poulin, E., Magoulas, A., Kornfield, I. & Ojeda, F.P. 2005. Origin, diversifi-
cation, and historical biogeography of the genus Trachurus (Perciformes: Carangidae). Molecular
Phylogenetics and Evolution 35, 496–507.
Carr, M.E., Strub, P.T., Thomas, A.C. & Blanco, J.L. 2002. Evolution of 1996–1999 La Niña and El Niño
conditions off the western coast of South America: a remote sensing perspective. Journal of Geophys-
ical Research-Oceans 107, C12, 3236 DOI:10.1029/2001JC001183.
Carrasco, F.D. 1997. Sublittoral macrobenthic fauna off Punta Coloso, Antofagasta, northern Chile: high
persistence of the polychaete assemblage. Bulletin of Marine Science 60, 443–459.
Carrasco, F.D., Gallardo, V.A. & Medrano, S. 1988. Sublittoral macrobenthic infaunal assemblages of two nearby
embayments from central Chile. Internationale Revue der Gesamten Hydrobiologie 73, 441–455.
Carrasco, S. & Santander, H. 1987. The El-Niño event and its influence on the zooplankton off Peru. Journal
of Geophysical Research-Oceans 92, 14,405–14,410.
Castilla, J.C. 1988. Earthquake-caused coastal uplift and its effects on rocky intertidal kelp communities.
Science 242, 440–443.
Castilla, J.C. 1996. The future Chilean marine park and preserves network and the concepts of conservation,
preservation and management according to the national legislation. Revista Chilena de Historia
Natural 69, 253–270.
Castilla, J.C. 1997. Chilean resources of benthic invertebrates: fishery, collapses, stock rebuilding and the role
of coastal management areas and national parks. In Developing and Sustaining World Fisheries
Resources. The State of Science and Management. Second World Fisheries Congress, D.A. Hancock
et al. (eds). Collingwood, Australia: Csiro Publishing, 130–135.
309
MARTIN THIEL ET AL.
Castilla, J.C. 1999. Coastal marine communities: trends and perspectives from human-exclusion experiments.
Castilla, J.C. 2000. Roles of experimental marine ecology in coastal management and conservation. Journal
Castilla, J.C. & Camus, P.A. 1992. The Humboldt–El Niño scenario — coastal benthic resources and anthro-
pogenic influences, with particular reference to the 1982/83 ENSO. South African Journal of Marine
Science 12, 703–712.
Castilla, J.C., Collins, A.G., Meyer, C.P., Guiñez, R. & Lindberg, D.R. 2002b. Recent introduction of the
dominant tunicate, Pyura praeputialis (Urochordata, Pyuridae) to Antofagasta, Chile. Molecular
Ecology 11, 1579–1584.
Castilla, J.C. & Defeo, O. 2005. Paradigm shifts needed for world fisheries. Science 309, 1324–1325.
Castilla, J.C. & Durán, L.R. 1985. Human exclusion from the rocky intertidal zone of central Chile: the effects
on Concholepas concholepas (Gastropoda). Oikos 45, 391–399.
Castilla, J.C. & Fernández, M. 1998. Small-scale benthic fisheries in Chile: on co-management and sustainable
use of benthic invertebrates. Ecological Applications 8, S124–S132.
Castilla, J.C., Fernández, M., Acuña, E., Bahamonde, N., Buschmann, A.H., Navarrete, S., Ulloa, O. & Yáñez,
E. 2005b. Ciencias del Mar. In Análisis y Proyecciones de la Ciencia Chilena 2005, J.E. Allende et al.
(eds). Santiago, Chile: Academia Chilena de Ciencias, 375–402.
Castilla, J.C. & Gelcich, S. 2006. Chile: Experience with management and exploitation areas for coastal
fisheries as building blocks for large-scale marine management. In Scaling Up Marine Management:
The Role of Marine Protected Areas. The World Bank Environment Department, Sustainable Devel-
opment Network. Report No. 36635-GLB, 45–57.
Castilla, J.C., Gelcich, S. & Defeo, O. 2007. Successes, lessons, and projections from experience in marine
benthic invertebrate artisanal fisheries in Chile (Chapter 2). In Fisheries Management: Challenges
and Accomplishment, M. McClanahan & J.C. Castilla (eds). Oxford, U.K.: Blackwell Publishing,
25–42.
Castilla, J.C., Guiñez, R., Alvarado, J.L., Pacheco, C. & Varas, M. 2000. Distribution, population structure,
population biomass and morphological characteristics of the tunicate Pyura stolonifera in the Bay of
Antofagasta, Chile. Marine Ecology Pubblicazioni Della Stazione Zoologica Di Napoli I 21, 161–174.
Castilla, J.C., Guinéz, R., Caro, A.U. & Ortíz, V. 2004a. Invasion of a rocky intertidal shore by the tunicate
Pyura praeputialis in the Bay of Antofagasta, Chile. Proceedings of the National Academy of Sciences
USA 101, 8517–8524.
Castilla, J.C., Lagos, N.A. & Cerda, M. 2004b. Marine ecosystem engineering by the alien ascidian Pyura
praeputialis on a mid-intertidal rocky shore. Marine Ecology Progress Series 268, 119–130.
Castilla, J.C., Lagos, N.A., Guiñez, R. & Largier, J. 2002a. Embayments and nearshore retention of plankton:
the Antofagasta Bay and other examples. In The Oceanography and Ecology of the Nearshore and
Bays in Chile, J.C. Castilla & J.L. Largier (eds). Santiago, Chile: Ediciones Universidad Católica de
Chile, 179–203.
Castilla, J.C. & Largier, J. (eds) 2002. The Oceanography and Ecology of the Nearshore and Bays in Chile.
Santiago, Chile: Ediciones Universidad Católica de Chile.
Castilla, J.C. & Neill, P.E. 2007. Marine bioinvasions in the southeastern Pacific: status, ecology, economic
impacts, conservation and management. In Marine Bioinvasions: Ecology, Conservation and Man-
agement Perspectives, G. Rilov & J. Crooks (eds). Berlin: Springer-Verlag. (in press).
Castilla, J.C. & Paine, R.T. 1987. Predation and community organization on eastern Pacific, temperate zone,
rocky intertidal shores. Revista Chilena de Historia Natural 60, 131–151.
Castilla, J.C., Uribe, M., Bahamonde, N., Clarke, M., Desqueyroux-Faúndez, R., Kong, I., Moyano, H.,
Rozbaczylo, N., Santelices, B., Valdovinos, C. & Zavala, P. 2005a. Down under the southeastern
Pacific: marine non-indigenous species in Chile. Biological Invasions 7, 213–232.
Castillo, J.G. 1968. Contribución al conocimiento de los Ofiuroideos Chilenos. Gayana (Chile) 14, 3–63.
Castro, L.R. 2001. Environmental conditions and larval survival during the winter spawning season of the
southernmost anchoveta stock off Chile. GLOBEC Newsletter 7, 15–17.
Castro, L.R., Bernal, P.A. & Troncoso, V.A. 1993. Coastal intrusion of copepods — mechanisms and conse-
quences on the population biology of Rhincalanus nasutus. Journal of Plankton Research 15, 501–515.
310
Castro, L.R. & Hernández, E.H. 2000. Early life survival of the anchoveta Engraulis ringens off central Chile
during the 1995 and 1996 winter spawning seasons. Transactions of the American Fisheries Society
129, 1107–1117.
Castro, L.R., Llanos, A., Blanco, J.L., Tarifeño, E., Escribano, R. & Landaeta, M.F. 2002. Latitudinal variations
in spawning habitat characteristics: influence on the early life history traits of the anchoveta Engraulis
ringens, off northern and central Chile. GLOBEC Newsletter 16, 42–45.
Castro, L.R., Salinas, G.R. & Hernández, E.H. 2000. Environmental influences on winter spawning of the
anchoveta Engraulis ringens off central Chile. Marine Ecology Progress Series 197, 247–258.
Castro, S.A., Camousseight, A., Munoz-Schick, M. & Jaksic, F.M. 2006. Rodulfo Amando Philippi, a naturalist
of major importance in the taxonomic classification of the biological diversity of Chile. Revista Chilena
de Historia Natural 79, 133–143.
Cerda, M. & Castilla, J.C. 2001. Diversity and biomass of macroinvertebrates in intertidal matrices of the
tunicate Pyura praeputialis (Heller, 1878) in the Bay of Antofagasta, Chile. Revista Chilena de Historia
Natural 74, 841–853.
Chaigneau, A. & Pizarro, O. 2005. Mean surface circulation and mesoscale turbulent flow characteristics in
the eastern South Pacific from satellite tracked drifters. Journal of Geophysical Research-Oceans 110,
C05014, DOI:10.1029/2004JC002628.
Chavez, F.P., Ryan, J., Lluch-Cota, S.E. & Niquen, M. 2003. From anchovies to sardines and back: multidecadal
change in the Pacific Ocean. Science 299, 217–221.
Chong, J., Oyarzún, C., Galleguillos, R., Tarifeño, E., Sepúlveda, R. & Ibáñez, C. 2005. Parametros biologico-
pesqueros de la jibia, Dosidicus gigas (Orbigny, 1835) (Cephalopoda: Ommastrephidae), frente a la
costa de chile central (29°S-40°S) durante 1993–1994. Gayana 69, 319–328.
Chopin, T., Buschmann, A.H., Halling, C., Troell, M., Kautsky, N., Neori, A., Kraemer, G.P., Zertuche-
González, J.A., Yarish, C. & Neefus, C. 2001. Integrating seaweeds into marine aquaculture systems:
a key toward sustainability. Journal of Phycology 37, 975–986.
Clarke, A. 1987. Temperature, latitude and reproductive effort. Marine Ecology Progress Series 38, 89–99.
Clarke, M., Ortiz, V. & Castilla, J.C. 1999. Does early development of the chilean tunicate Pyura praeputialis
(Heller, 1878) explain the restricted distribution of the species? Bulletin of Marine Science 65, 745–754.
Coale, K.H., Johnson, K.S., Fitzwater, S.E., Blain, S.P., Stanton, T.P. & Coley, T.L. 1998. Iron ex-I, an in situ
iron enrichment experiment: experimental design, implementation and results. Deep-Sea Research II
45, 919–945.
Codispoti, L.A. & Christensen, J.P. 1985. Nitrification, denitrification and nitrous oxide cycling in the eastern
tropical South Pacific ocean. Marine Chemistry 16, 277–300.
Codispoti, L.A., Friederich, G.E., Packard, T.T., Glover, H.E., Kelly, P.J., Spinrad, R.W., Barber, R.T., Elkins,
J.W., Ward, B.B., Lipschultz, F. & Lostaunau, N. 1986. High nitrite levels off northern Perú: a signal
of instability in the marine denitrification rate. Science 233, 1200–1202.
Cohen, C.S. & Strathmann, R.R. 1996. Embryos at the edge of tolerance: effects of environment and structure
of egg masses on supply of oxygen to embryos. Biological Bulletin 190, 8–15.
Collantes, G., Merino, A. & Lagos, V. 2002. Fenología de la gametogénesis, madurez de conceptáculos,
fertilidad y embriogénesis en Durvillea antarctica (Chamisso) Hariot (Phaeophyta, Durvillaeales).
Revista de Biología Marina y Oceanografia (Chile) 37, 83–112.
Coloma, C., Marchant, M. & Hebbeln, D. 2005. Foraminíferos planctónicos durante El Niño 1997–1998 del
área de Coquimbo (30°S; 73°W), Chile. Gayana 69, 48–77.
Colombini, I., Aloia, A., Fallaci, M., Pezzoli, G. & Chelazzi, L. 2000. Temporal and spatial use of stranded
wrack by the macrofauna of a tropical sandy beach. Marine Biology 136, 531–541.
CONAMA-PNUD. 2006. Conservación de la biodiversidad de importancia mundial a lo largo de la costa
Chilena. Areas Marinas y Costeras Protegidas de Múltiples Usos. Isla Grande de Atacama, Lafken
Mapu Lahual, Francisco Coloane. Ocho Libros Editores, Santiago, Chile.
Conkright, M.E., Locarnini, R.A., Garcia, H.E., O’Brien, T.D., Boyer, T.P., Stephensons, C. & Antonov, J.I.
2002. World Ocean Atlas 2001: objective analyses, data statistics, and figures. Technical report,
National Oceanographic Data Center. Silver Spring, MD, USA.
Connolly, S.R., Menge, B.A. & Roughgarden, J. 2001. A latitudinal gradient in recruitment of intertidal
invertebrates in the northeast Pacific Ocean. Ecology 82, 1799–1813.
311
MARTIN THIEL ET AL.
Contreras, H., Defeo, O. & Jaramillo, E. 1999. Life history of Emerita analoga (Stimpson) (Anomura,
Hippidae) in a sandy beach of south central Chile. Estuarine Coastal and Shelf Science 48, 101–112.
Contreras, H. & Jaramillo, E. 2003. Geographical variation in natural history of the sandy beach isopod
Excirolana hirsuticauda Menzies (Cirolanidae) on the Chilean coast. Estuarine Coastal and Shelf
Science 58, 117–126.
Contreras, H., Jaramillo, E. & Quijón, P. 2000. Natural history of Emerita analoga (Stimpson) (Anomura,
Hippidae) in a sandy beach of northern Chile. Revista Chilena de Historia Natural 73, 705–715.
Cornejo, M., Farias, L. & Paulmier, A. 2006. Temporal variability in N2O water content and its air-sea exchange
in an upwelling area off central Chile (36°S). Marine Chemistry 101, 85–94.
Correa, J.A., Avila, M. & Santelices, B. 1985. Effects of some environmental factors on growth of sporelings
of two species of Gelidium (Rodophyta). Aquaculture 44, 221–227.
Correa, J.A., Castilla, J.C., Ramírez, M., Varas, M., Lagos, N.A., Vergara, S., Moenne, A., Roman, D. &
Brown, M.T. 1999. Copper, copper mine tailings and their effect on marine algae in northern Chile.
Journal of Applied Phycology 11, 57–67.
Cowie, G.L., Calvert, S.E., Pedersen, T.F., Schulz, H. & von Rad, U. 1999. Organic content and preservational
controls in surficial shelf and slope sediments from the Arabian Sea (Pakistan margin). Marine Geology
161, 23–38.
Crisp, D.J., Yule, A.B. & White, K.N. 1985. Feeding by oyster larvae the functional response, energy budget
and a comparison with mussel larvae. Journal of the Marine Biological Association of the United
Kingdom 65, 759–784.
Cubillos, L.A., Arcos, D.F., Bucarey, D.A. & Canales, M.T. 2001. Seasonal growth of small pelagic fish off
Talcahuano, Chile (37°S, 73°W): a consequence of their reproductive strategy to seasonal upwelling?
Aquatic Living Resources 14, 115124.
Cubillos, L.A., Bucarey, D.A. & Canales, M.T. 2002. Monthly abundance estimation for common sardine
Strangomera bentincki and anchovy Engraulis ringens in the central-southern area off Chile (34–40°S).
Fisheries Research 57, 117–130.
Cuevas, L.A., Daneri, G., Jacob, B. & Montero, P. 2004. Microbial abundance and activity in the seasonal
upwelling area off Concepción (36°S), central Chile: a comparison of upwelling and non-upwelling
conditions. Deep-Sea Research II 51, 2427–2440.
Cury, P., Bakun, A., Crawford, R.J.M., Jarre, A., Quinones, R.A., Shannon, L.J. & Verheye, H.M. 2000. Small
pelagics in upwelling systems: patterns of interaction and structural changes in ‘wasp-waist’ ecosys-
tems. ICES Journal of Marine Science 57, 603–618.
Cury, P. & Roy, C. 1989. Optimal environmental window and pelagic fish recruitment success in upwelling
areas. Canadian Journal of Fisheries and Aquatic Sciences 46, 670–680.
Cushing, D.H. 1971. Upwelling and the production of fish. Advances in Marine Biology 9, 255–334.
Dahlhoff, E. & Somero, G.N. 1993. Effects of temperature on mitochondria from abalone (Genus Haliotis)
adaptive plasticity and its limits. Journal of Experimental Biology 185, 151–168.
Dall, W.H. 1909. Report on a collection of shells from Peru, with a summary of the littoral marine Mollusca
of the Peruvian zoological province. Proceedings of the United States National Museum 37, 147–293
plus 9 plates.
Daneri, G., Dellarossa, V., Quiñones, R., Jacob, B., Montero, P. & Ulloa, O. 2000. Primary production and
community respiration in the Humboldt Current System off Chile and associated oceanic areas. Marine
Dantagnan, P. 1993. Fertilidad y efecto de factores ambientales sobre el desarrollo inicial de las fases
isomórficas de Gelidium chilense (Montagne) Santelices et Montalvo. Honors thesis, Universidad
Católica del Norte, Coquimbo, Chile.
Darwin, C.R. 1851. A Monograph on the Subclass Cirripedia. The Lepadidae; or, Pedunculated Cirripedes.
London: Ray Society.
Darwin, C.R. 1854. A Monograph on the Subclass Cirripedia 2. The Balanidae, Verrucidae, etc. London: Ray
Society.
Da Silva, A.M., Young, C.C. & Levitus, S. 1994. Atlas of Surface Marine Data. Silver Spring, Maryland: U.S.
Departament of Commerce, NOAA.
312
Dayton, P.K. 1985. The structure and regulation of some South American kelp communities. Ecological
Dayton, P.K., Currie, V., Gerrodette, T., Keller, B.D., Rosenthal, R. & Ven Tresca, D. 1984. Path dynamics
and stability of some Californian kelp communities. Ecological Monographs 54, 253–289.
Defeo, O. & Castilla, J.C. 2005. More than one bag for the world fishery crisis and keys for co-management
successes in selected artisanal Latin American shellfisheries. Reviews in Fish Biology and Fisheries
15, 265–283.
Defeo, O., McClanahan, T. & Castilla, J.C. 2007. A brief history of fisheries management with emphasis on
societal participatory roles. In Fisheries Management: Challenges and Accomplishment,
T. McClanahan & J.C. Castilla (eds). Oxford, U.K.: Blackwell Publishing, 3–21.
Defeo, O. & McLachlan, A. 2005. Patterns, processes and regulatory mechanisms in sandy beach macrofauna:
a multi-scale analysis. Marine Ecology Progress Series 295, 1–20.
Delille, D., Marty, G., CansemiSoullard, M. & Frankignoulle, M. 1997. Influence of subantarctic Macrocystis
bed metabolism in diel changes of marine bacterioplankton and CO2 fluxes. Journal of Plankton
Research 19, 1251–1264.
Dell, R.K. 1971. The marine Mollusca of the Royal Society expedition to southern Chile. Records of the
Dominion Museum 7, 155–233.
Dellinger, T. & Trillmich, F. 1999. Fish prey of the sympatric Galapagos fur seals and sea lions: seasonal
variation and niche separation. Canadian Journal of Zoology 77, 1204–1216.
Desqueyroux, R. & Moyano, H. 1987. Zoogeografía de demospongias chilenas. Boletín de la Sociedad de
Biología de Concepción (Chile) 58, 39–66.
DiBacco, C. & Levin, L.A. 2000. Development and application of elemental fingerprinting to track the dispersal
of marine invertebrate larvae. Limnology and Oceanography 45, 871–880.
Duarte, C., Jaramillo, E. & Contreras, H. 2004. Zonación de Orchestoidea tuberculata (Nicolet) en playas de
arena en la costa de Valdivia. Libro Resumen XXIV Congreso de Ciencias del Mar, Coquimbo, Chile, 233.
Duarte, W.E., Asencio, G. & Moreno, C.A. 1996. Long-term changes in population density of Fissurella picta
and Fissurella limbata (Gastropoda) in the marine reserve of Mehuin, Chile. Revista Chilena de
Historia Natural 69, 45–56.
Ducklow, H.W. 2000. Bacterial production and biomass in the oceans. In Microbial Ecology of the Oceans,
D. Kirchman (ed.). New York: Wiley-Liss, 85–120.
Duffy, D., Hays, C. & Plenge, M. 1984. The conservation status of Peruvian seabirds. In Status and Conservation
of the World’s Seabirds, J.P. Croxall et al. (eds). Cambridge, U.K.: ICBP Technical Publication, 245–259.
Dugan, J.E., Hubbard, D.M., McCrary, M.D. & Pierson, M.O. 2003. The response of macrofauna communities
and shorebirds to macrophyte wrack subsidies on exposed sandy beaches of southern California.
Dugan, J.E., Jaramillo, E., Hubbard, D.M., Contreras, H. & Duarte, C. 2004. Competitive interactions in
macroinfaunal animals of exposed sandy beaches. Oecologia 139, 630–640.
Dunton, K., Goodall, J., Schonberg, S., Grebmeier, J. & Maidment, D. 2005. Multi-decadal synthesis of
benthic–pelagic coupling in the western arctic: role of cross-shelf advective processes. Deep-Sea
Research II 52, 3462–3477.
Durán, L.R. & Castilla, J.C. 1989. Variation and persistence of the middle rocky intertidal community of
central Chile, with and without human harvesting. Marine Biology 103, 555–562.
Ebensperger, L.A. & Botto-Mahan, C. 1997. Use of habitat, size of prey, and food-niche relationships of two
sympatric otters in southernmost Chile. Journal of Mammalogy 78, 429–434.
Ebensperger, L.A. & Castilla, J.C. 1992. Habitat selection in land by the marine otter, Lutra felina, at Pan de
Azúcar island, Chile. Revista Chilena de Historia Natural 65, 429–434.
Edding, M. & Blanco, M. 2001. Estrategias de vigilancia biológica del evento El Niño. In El Niño en América
Latina, impactos biológicos y sociales, J. Tarazona et al. (eds). Lima: Consejo Nacional de Ciencia
y Tecnología, 127–136.
Edding, M., Fonck, E.A. & Macchiavello, J.E. 1994. Lessonia. In Biology of Economic Algae, I. Akatsuka
(ed.). The Hague: The Hague Academic Publishing, 407–446.
Edding, M., Fonck, E.A., Orrego, P., Venegas, M. & Macchiavello, J. 1993. A comparison between two populations
of Lessonia trabeculata (Phaeophyta, Laminariales) microscopic stages. Hydrobiologia 261, 231–237.
313
MARTIN THIEL ET AL.
Edding, M. & Tala, F. 2003. Development of techniques for the cultivcation of Lessonia trabeculata Villouta
et Santelices (Phaeophyceae, Laminariales) in Chile. Aquaculture Research 34, 507–515.
Edding, M., Venegas, M., Orrego, P. & Fonck, E.A. 1990. Culture and growth of Lessonia trabeculata
(Phaeophyta, Laminariales) juvenile sporophytes in La Herradura de Guayacán Bay, northern Chile.
Hydrobiologia 204, 361–366.
Edwards, M.S. 2004. Estimating scale-dependency in disturbance impacts: El Niño and giant kelp forests in
Eissler, Y. & Quiñones, R.A. 1999. Microplanktonic respiration off northern Chile during El Niño 1997–1998.
Journal of Plankton Research 21, 2263–2283.
Ekman, S. 1953. Zoogeography of the Sea. London: Sidgwick & Jackson.
Ellenberg, U., Mattern, T., Seddon, P. & Luna-Jorquera, G. 2006. Physiological and reproductive consequences
of human disturbance in Humboldt penguins: the need for species-specific visitor management.
Biological Conservation 133, 95–106.
Ellis, J.C., Farina, J.M. & Witman, J.D. 2006. Nutrient transfer from sea to land: the case of gulls and
cormorants in the Gulf of Maine. Journal of Animal Ecology 75, 565–574.
Enfield, D.B., Cornejorodriguez, M.D., Smith, R.L. & Newberger, P.A. 1987. The Equatorial source of
propagating variability along the Peru coast during the 1982–1983 El Niño. Journal of Geophysical
Research-Oceans 92, 14,335–14,346.
Escribano, R. 1998. Population dynamics of Calanus chilensis in the Chilean Eastern Boundary Humboldt
Current. Fisheries Oceanography 7, 245–251.
Escribano, R., Daneri, G., Farias, L., Gallardo, V.A., González, H.E., Gutiérrez, D., Lange, C.B., Morales,
C.E., Pizarro, O., Ulloa, O. & Braun, M. 2004a. Biological and chemical consequences of the 1997–1998
El Niño in the Chilean coastal upwelling system: a synthesis. Deep-Sea Research II 51, 2389–2411.
Escribano, R., Fernández, M. & Aranís, A. 2003. Physical-chemical processes and patterns of diversity of the
Chilean eastern boundary pelagic and benthic marine ecosystems: an overview. Gayana 67, 190–205.
Escribano, R. & Hidalgo, P. 2000a. Spatial distribution of copepods in the north of the Humboldt Current
region off Chile during coastal upwelling. Journal of the Marine Biological Association of the United
Kingdom 80, 283–290.
Escribano, R. & Hidalgo, P. 2000b. Influence of El Niño and La Niña on the population dynamics of Calanus
chilensis in the Humboldt Current ecosystem of northern Chile. Ices Journal of Marine Science 57,
1867–1874.
Escribano, R., Hidalgo, P. & Krautz, C. Zooplankton associated with the oxygen minimum zone off the
northern upwelling region of Chile in March 2000. Deep Sea Research Part II. The Oceanography
of the eastern South Pacific (accepted).
Escribano, R., Hidalgo, P., González, H.E., Araneda, A., Giesecke, R., & Manríquez, K. Interannual and
seasonal variability of metazooplankton in the central/south upwelling region off Chile. (submitted)
Progress in Oceanography, Special Issue ‘The Structure and Functioning of the Coastal Upwelling
System Off Central/South Chile’ R. Escribano & W. Schneider (eds).
Escribano, R., Marín, V.H. & Hidalgo, P. 2001. The influence of coastal upwelling on the distribution of
Calanus chilensis in the Mejillones Peninsula (northern Chile): implications for its population dynam-
ics. Hydrobiologia 453, 143–151.
Escribano, R., Marín, V.H., Hidalgo, P. & Olivares, G. 2002. Physical-biological interactions in the pelagic
ecosystem of the nearshore zone of the northern Humboldt Current System. In The Oceanography
and Ecology of the Nearshore and Bays in Chile, J.C. Castilla & J.L. Largier (eds). Santiago, Chile:
Ediciones Universidad Católica de Chile, 145–175.
Escribano, R., Marín, V.H. & Irribarren, C. 2000. Distribution of Euphausia mucronata at the upwelling area
of Peninsula Mejillones, northern Chile: the influence of the oxygen minimum layer. Scientia Marina
64, 69–77.
Escribano, R. & McLaren, I. 1999. Production of Calanus chilensis in the upwelling area of Antofagasta,
northern Chile. Marine Ecology Progress Series 177, 147–156.
Escribano, R. & Rodríguez, L. 1994. Life cycle of Calanus chilensis Brodsky in Bay of San Jorge, Antofagasta,
Chile. Hydrobiologia 293, 289–294.
314
Escribano, R., Rosales, S.A. & Blanco, J.L. 2004b. Understanding upwelling circulation off Antofagasta
(northern Chile): a 3-dimensional numerical-modeling approach. Continental Shelf Research 24, 37–53.
Espinoza, F. 2006. Patrones reproductivos de Acanthocyclus hassleri Rathbun 1898 and Acanthocyclus gayi
Milne-Edwards & Lucas 1844 en Chile central. Honors thesis, Universidad de Valparaíso, Valparaíso,
Chile.
FAL. 1991. Fishery and Aquaculture Law. Ministerio de Economía Fomento y Reconstrucción, Gobierno de
Chile. Ley No. 18892, D.S. No. 430.
FAO. 1995. Code of Conduct for Responsible Fisheries. Rome: FAO.
FAO. 2003. Fisheries Technical Paper n°433. The ecosystem approach to fisheries. Issues, terminology,
principles, institutional foundations, implementation and outlook. Rome: FAO.
FAO. 2006. Directory of Chilean Aquaculture and Fisheries. Santiago, Chile: Technopress S.A., thirteenth edition.
Farías, L., Graco, M. & Ulloa, O. 2004. Temporal variability of nitrogen cycling in continental-shelf sediments
of the upwelling ecosystem off central Chile. Deep-Sea Research II 51, 2491–2505.
Fariña, J.M., Castilla, J.C. & Ojeda, F.P. 2003a. The ‘idiosyncratic’ effect of a ‘sentinel’ species on contam-
inated rocky intertidal communities. Ecological Applications 13, 1533–1552.
Fariña, J.M., Palma, A.T. & Ojeda, F.P. A food web perspective of the subtidal communities off the temperate
Chilean coast. In Food Webs and the Dynamics of Marine Benthic Ecosystems, T.R. McClanahan &
G.M. Branch (eds). Oxford: Oxford University Press (in press).
Fariña, J.M., Salazar, S., Wallem, K.P., Witman, J.D. & Ellis, J.C. 2003b. Nutrient exchanges between marine
and terrestrial ecosystems: the case of the Galapagos sea lion Zalophus wollebaecki. Journal of Animal
Ecology 72, 873–887.
Farrell, T.M., Bracher, D. & Roughgarden, J. 1991. Cross-shelf transport causes recruitment to intertidal
populations in central California. Limnology and Oceanography 36, 279–288.
Faugeron, S., Martínez, E.A., Correa, J.A. & Billot, C. 2005. Long-term copper mine waste disposal in northern
Chile associated with gene flow disruption of the intertidal kelp Lessonia nigrescens. Marine Ecology
Faugeron, S., Valero, M., Destombe, C., Martinez, E.A. & Correa, J.A. 2001. Hierarchical spatial structure
and discriminant analysis of genetic diversity in the red alga Mazzaella laminarioides (Gigartinales,
Rhodophyta). Journal of Phycology 37, 705–716.
Fernández, D., Escribano, R. & Hidalgo, P. 2002. Distribución de eufáusidos en el sistema de surgencia frente
a la península de Mejillones (23°S) asociada a condiciones previas durante El Niño 1997/98. Inves-
tigaciones Marinas, Valparaíso 31, 56–68.
Fernández, E., Córdova, C. & Tarazona, J. 1999. Condiciones del bosque submareal de Lessonia trabeculata
en la isla Independencia durante el evento El Niño 1997–1998. Revista Peruana de Biología Volumen
extraordinario, 47–59.
Fernández, M. & Brante, A. 2003. Brood care in Brachyuran crabs: the effect of oxygen provision on
reproductive costs. Revista Chilena de Historia Natural 76, 157–168.
Fernández, M., Calderon, R., Cifuentes, M. & Pappalardo, P. 2006b. Brooding behaviour and cost of brooding
in small body size brachyuran crabs. Marine Ecology Progress Series 309, 213–220.
Fernández, M. & Castilla, J.C. 2000. Recruitment of Homalaspis plana in intertidal habitats of central Chile
and implications for the current use of management and Marine Protected Areas. Marine Ecology
Fernández, M. & Castilla, J.C. 2005. Marine conservation in Chile: historical perspective, lessons, and
challenges. Conservation Biology 19, 1752–1762.
Fernández, M., González, C. & Jeno, K. 2007. Large-scale spatial patterns of gonad production in Perumytilus
purpuratus across a transition zone in central Chile. Journal of Shellfish Research (in press).
Fernández, M., Jaramillo, E., Marquet, P.A., Moreno, C.A., Navarrete, S.A., Ojeda, F.P., Valdovinos, C.R. &
Vásquez, J.A. 2000. Diversity, dynamics and biogeography of Chilean benthic nearshore ecosystems:
an overview and guidelines for conservation. Revista Chilena de Historia Natural 73, 797–830.
Fernández, M., Pappalardo, P. & Jeno, K. 2006a. The effects of temperature and oxygen availability on
intracapsular development of Acanthina monodon (Gastropoda: Muricidae). Revista Chilena de His-
toria Natural 79, 155–167.
315
MARTIN THIEL ET AL.
Fernández, M., Pardo, L.M. & Baeza, J.A. 2002. Patterns of oxygen supply in embryo masses of brachyuran
crabs throughout development: the effect of oxygen availability and chemical cues in determining
female brooding behavior. Marine Ecology Progress Series 245, 181–190.
Fernández, M., Ruiz-Tagle, N., Cifuentes, S., Pörtner, H.O. & Arntz, W. 2003. Oxygen-dependent asynchrony
of embryonic development in embryo masses of brachyuran crabs. Marine Biology 142, 559–565.
Fiedler, P.C. & Lavín, M.F. 2006. Introduction: a review of eastern tropical Pacific oceanography. Progress
in Oceanography 69, 94–100.
Figueroa, D. 2002. Forcing of physical exchanges in the nearshore Chilean ocean. In The Oceanography and
Ecology of the Nearshore and Bays in Chile, J.C. Castilla & J.L. Largier (eds). Santiago, Chile:
Figueroa, D. & Moffat, C. 2000. On the influence of topography in the induction of coastal upwelling along
the Chilean coast. Geophysical Research Letters 27, 3905–3908.
Fonseca, T. 1989. An overview of the poleward undercurrent and upwelling along the Chilean coast. In
Poleward Flows Along Eastern Ocean Boundaries, S.J. Neshyba et al. (eds). New York: Springer-
Verlag, 203–218.
Fonseca, T. & Farías, M. 1987. Estudio del proceso de surgencia en la costa chilena utilizando percepción
remota. Investigaciones Pesqueras 34, 33–46.
Forward, R.B., Tankersley, R.A. & Reinsel, K.A. 1998. Selective tidal stream transport of spot (Leistomus
xanthurus, Lacepede) and pinfish (Lagodon rhomboides, Linnaeus) larvae: contribution of circatidal
rhythms in activity. Journal of Experimental Marine Biology and Ecology 226, 19–32.
Fossing, H., Gallardo, V.A., Jorgensen, B.B., Hüttel, M., Nielsen, L.P., Schulz, H., Canfield, D.E., Forster, S.,
Glud, R.N., Gundersen, J.K., Kuver, J., Ramsing, N.B., Teske, A., Thamdrup, B. & Ulloa, O. 1995.
Concentration and transport of nitrate by the mat-forming sulfur bacterium Thioploca. Nature 374,
713–715.
Frederich, M. & Pörtner, H.O. 2000. Oxygen limitation of thermal tolerance defined by cardiac and ventilatory
performance in spider crab, Maja squinado. American Journal of Physiology-Regulatory Integrative
and Comparative Physiology 279, R1531–R1538.
Freeland, H.J., Gatien, G., Huyer, A. & Smith, R.L. 2003. Cold halocline in the northern California Current:
an invasion of subarctic water. Geophysical Research Letters 30, 3, 1141, DOI:10.1029/2002GL016663.
Frere, E., Gandini, P., Ruiz, J. & Vilina, Y.A. 2004. Current status and breeding distribution of red-legged cormorant
Phalacrocorax gaimardi along the Chilean coast. Bird Conservation International 14, 113–121.
Frid, C.L., Buchanan, J.B. & Garwood, P.R. 1996. Variability and stability in benthos: twenty-two years of
monitoring off Northumberland. ICES Journal of Marine Science 53, 978–980.
Fuenzalida, R. 1990. Variabilidad temporal de un índice de surgencia para la zona de Iquique (Lat. 20°S).
Investigación Científica y Tecnológica, Serie Ciencias Marinas 1, 37–47.
Fuenzalida, R., Schneider, W., Garcés-Vargas, J., Bravo, L. & Lange, C.B. Accepted. Vertical and horizontal
extension of the oxygen minimum zone in the eastern South Pacific Ocean. Deep Sea Research II.
Furness, R. & Monaghan, P. 1987. Seabird Ecology. Glasgow: Blackie.
Gajardo, G., Cancino, J.M. & Navarro, J.M. 2002. Genetic variation and population structure in the marine
snail Chorus giganteus (Gastropoda: Muricidae), an overexploited endemic resource from Chile.
Gallardo, V.A. 1963. Notas sobre la densidad de la fauna bentónica en el sublitoral del norte de Chile. Gayana
Oceanológica 10, 3–15.
Gallardo, V.A. 1977. Large benthic microbial communities in sulfide biota under Perú-Chile subsurface
countercurrent. Nature 268, 331–332.
Gallardo, V.A., Cañete, J.I., Roa, R., Enriquez-Briones, S. & Baltazar, M. 1994. Recruitment of the squat
lobster Pleuroncodes monodon on the continental shelf off central Chile. Journal of Crustacean
Biology 14, 665–669.
Gallardo, V.A., Palma, M., Carrasco, F.D., Gutiérrez, D., Levin, L.A. & Cañete, J.I. 2004. Macrobenthic
zonation caused by the oxygen minimum zone on the shelf and slope off central Chile. Deep-Sea
Research II 51, 2475–2490.
Galleguillos, R., Troncoso, L., Oyarzún, C., Astorga, M. & Peñaloza, M. 2000. Genetic differentiation in
Chilean hake Merluccius gayi gayi (Pisces: Merlucciidae). Hydrobiologia 420, 49–54.
316
García, J.C. 1996. Fenología y ciclo de vida de Glossophora kunthii (C.A.) J. Agardh de poblaciones inter y
submareales en Puerto Aldea IV Región. Honors thesis, Universidad Católica del Norte, Coquimbo,
Chile.
Gelcich, S. 1999. Dimensión fractal de macroalgas submareales: su relación con la fauna epibionte. Honors
thesis, Universidad Católica del Norte, Coquimbo, Chile.
Gelcich, S., Edwards-Jones, G., Kaiser, M.J. & Castilla, J.C. 2006. Co-management policy can reduce
resilience in traditionally managed marine ecosystems. Ecosystems 9, 951–966.
Gelcich, S., Edwards-Jones, G., Kaiser, M. & Watson, E. 2005. Using discourses for policy evaluation: the
case of marine common property rights in Chile. Society and Natural Resources 18, 377–391.
Giesecke, R. & González, H.E. 2004. Feeding of Sagitta enflata and vertical distribution of chaetognaths in
relation to low oxygen concentrations. Journal of Plankton Research 26, 475–486.
Giraldo, A., Escribano, R. & Marín, V. 2002. Spatial distribution of Calanus chilensis off Mejillones Peninsula
(northern Chile): ecological consequences upon coastal upwelling. Marine Ecology Progress Series
230, 225–234.
Glud, R.N., Gundersen, J.K. & Holby, O. 1999. Benthic in situ respiration in the upwelling area off central
Chile. Marine Ecology Progress Series 186, 9–18.
Glynn, P.W. 1988. El Niño Southern Oscillation 1982–1983 — nearshore population, community, and eco-
system responses. Annual Review of Ecology and Systematics 19, 309–345.
Godoy, N.E. 2000. Macrocystis integrifolia (Laminariales, Phaeophyta) en el norte de Chile: distribución
espacial y fauna asociada. Honors thesis, Universidad Católica del Norte, Coquimbo, Chile.
Gómez, I., Figueroa, F.L., Huovinen, P., Ulloa, N. & Morales, V. 2005a. Photosynthesis of the red alga
Gracilaria chilensis under natural solar radiation in an estuary in southern Chile. Aquaculture 244,
369–382.
Gómez, I., Figueroa, F.L., Ulloa, N., Morales, V., Lovengreen, C., Huovinen, P. & Hess, S. 2004. Patterns of
photosynthesis in 18 species of intertidal macroalgae from southern Chile. Marine Ecology Progress
Series 270, 103–116.
Gómez, I., Ulloa, N. & Orostegui, M. 2005b. Morpho-functional patterns of photosynthesis and UV sensitivity
in the kelp Lessonia nigrescens (Laminariales, Phaeophyta). Marine Biology 148, 231–240.
Gómez, I. & Westermeier, R.C. 1991. Frond regrowth from basal disk in Iridaea laminarioides (Rhodophyta,
Gigartinales) at Mehuin, southern Chile. Marine Ecology Progress Series 73, 83–91.
Gomez-Uchida, D., Weetman, D., Hauser, L., Galleguillos, R. & Retamal, M. 2003. Allozyme and AFLP
analyses of genetic population structure in the hairy edible crab Cancer setosus from the Chilean
coast. Journal of Crustacean Biology 23, 486–494.
Gong, D.Y. & Wang, S.W. 1999. Definition of Antarctic oscillation index. Geophysical Research Letters 26,
459–462.
González, A. & Marín, V.H. 1998. Distribution and life cycle of Calanus chilensis and Centropages brachiatus
(Copepoda) in Chilean coastal waters: a GIS approach. Marine Ecology Progress Series 165, 109–117.
González, C. 1998. Efecto de marejadas asociadas al evento de El Niño 1997 sobre una pradera de Gracilaria
en la bahía de la Herradura de Guayacán (Coquimbo, IV Región). Libro Resumen XVIII Congreso
Ciencias del Mar. Iquique, Chile: Sociedad Ciencias del Mar, 161.
González, H.E., Bernal, P. & Ahumada, R. 1987. Desarrollo de dominancia local en la taxocenosis de
fitoplancton de Bahía de Concepción, Chile, durante un evento de surgencia. Revista Chilena de
González, H.E., Daneri, G., Figueroa, D., Iriarte, J.L., Lefevre, N., Pizarro, G., Quiñones, R., Sobarzo, M. &
Troncoso, A. 1998. Primary production and its fate in the pelagic food web and deep-sea, and ocean-
atmosphere CO2 exchange in the northern Humboldt Current (23°S): possible effects of the 1997–1998
El Niño in Chile. Revista Chilena de Historia Natural 71, 429–458.
González, H.E., Giesecke, R., Vargas, C.A., Pávez, A., Iriarte, J., Santibáñez, P., Castro, L., Escribano, R. &
Pages, F. 2004b. Carbon cycling through the pelagic foodweb in the northern Humboldt Current off
Chile (23°S). ICES Journal of Marine Science 61, 572–584.
González, H.E., Hebbeln, D., Iriarte, J. & Marchant, M. 2004a. Downward fluxes of faecal material and
microplankton at 2300 m depth in the oceanic area off Coquimbo (30°S), Chile, during 1993–1995.
Deep-Sea Research II 51, 2457–2474.
317
MARTIN THIEL ET AL.
González, H.E., Ortíz, V.C. & Sobarzo, M. 2000a. The role of faecal material in the particulate organic carbon
flux in the northern Humboldt Current, Chile (23°S), before and during the 1997–1998 El Niño.
González, H.E., Pages, F., Sobarzo, M. & Escribano, R. 2002. Effects of the 1997/98 El Niño on the
oceanographic conditions and zooplankton community structure in the coastal upwelling system off
northern Chile. Investigaciones Marinas, Valparaíso (extended abstract) 30, 112–114.
González, H.E., Pantoja, S., Iriarte, J. & Bernal, P.A. 1989. Winter-spring variability of size fractioned
autotrophic biomass in Concepción Bay, Chile. Journal of Plankton Research 11, 1157–1167.
González, H.E., Sobarzo, M., Figueroa, D. & Nothig, E.M. 2000b. Composition, biomass and potential grazing
impact of the crustacean and pelagic tunicates in the northern Humboldt Current area off Chile:
differences between El Niño and non-El Niño years. Marine Ecology Progress Series 195, 201–220.
González, J., Meneses, I. & Vásquez, J.A. 1997. Field studies in Chondracanthus chamissoi (C. Agardh)
Kützing. Seasonal and spatial variations in life cycle phases. Biología Pesquera (Chile) 26, 3–12.
González, J., Stotz, W., Garrido, J., Orensanz, J.M., Parma, A.M., Tapia, C. & Zuleta, A. 2006. The Chilean TURF
system: how is it performing in the case of the loco fishery? Bulletin of Marine Science 78, 499–527.
González, J., Tapia, C., Wilson, A., Stotz, W., Orensanz, J.M., Parma, A.M., Valero, J., Catrilao, M. & Garrido,
J. 2004. Bases biológicas para la evaluación y manejo de metapoblaciones de loco en las III y IV
Regiones. Project FIP No. 2002-16, Informe Final. Valparaíso, Chile: IFOP.
González, R.R. & Quiñones, R.A. 2000. Pyruvate oxidoreductases involved in glycolytic anaerobic metabolism
of polychaetes from the continental shelf off central-south Chile. Estuarine Coastal and Shelf Science
51, 507–519.
González, R.R. & Quiñones, R.A. 2002. Ldh activity in Euphausia mucronata and Calanus chilensis: impli-
cations for vertical migration behaviour. Journal of Plankton Research 24, 1349–1356.
González, S.A., Stotz, W. & Aguilar, M. 2001. Stranding of scallops related to epiphytic seaweeds on the
coast of northern Chile. Journal of Shellfish Research 20, 85–88.
Gooday, A.J., Bernhard, J.M., Levin, L.A. & Suhr, S.B. 2000. Foraminifera in the Arabian Sea oxygen
minimum zone and other oxygen-deficient settings: taxonomic composition, diversity, and relation to
metazoan faunas. Deep-Sea Research II 47, 25–54.
Gorshkov, S.G. 1985. World Ocean Atlas. Pacific Ocean. Oxford, U.K.: Pregamon Press.
Graco, M., Gutiérrez, D. & Farías, L. 2006. Inter-annual variability of the pelagic-benthic coupling in the
upwelling system off central Chile. Advances in Geosciences 6, 127–132.
Graf, G. 1989. Benthic-pelagic coupling in a deep-sea benthic community. Nature 341, 437–439.
Graham, M.H. 2004. Effects of local deforestation on the diversity and structure of Southern California giant
kelp forest food webs. Ecosystems 7, 341–357.
Graham, M.H., Vásquez, J.A. & Buschmann, A.H. 2007. Global ecology of the giant kelp Macrocystis: from
ecotypes to ecosystems. Oceanography and Marine Biology: An Annual Review 45, 39–88.
Grantham, B.A., Chan, F., Nielsen, K.J., Fox, D.S., Barth, J.A., Huyer, A., Lubchenco, J. & Menge, B.A.
2004. Upwelling-driven nearshore hypoxia signals ecosystem and oceanographic changes in the
northeast Pacific. Nature 429, 749–754.
Graves, J.E. & Somero, G.N. 1982. Electrophoretic and functional enzymatic evolution in four species of
eastern Pacific barracudas from different thermal habitats. Evolution 36, 97–106.
Grünewald, A., Morales, C., González, H.E., Sylvester, C. & Castro, L. 2002. Grazing impact of copepod
assemblages and gravitational flux in coastal and oceanic waters off central Chile during two con-
trasting seasons. Journal of Plankton Research 24, 55–68.
Guerra, C., Fitzpatrick, L., Aguilar, R. & Luna-Jorquera, G. 1988. Location and characterization of new nesting
sites for Gray gulls, Larus modestus, in the Atacama desert, northern Chile. Le Gerfaut 78, 121–129.
Guerra, C., van Waerebeek, K., Porflitt, G. & Luna-Jorquera, G. 1987. Presencia de cetáceos frente a la segunda
región de Chile. Estudios Oceanológicos (Chile) 6, 87–97.
Guizien, K., Brochier, T., Duchene, J.C., Koh, B.S. & Marsaleix, P. 2006. Dispersal of Owenia fusiformis
larvae by wind-driven currents: turbulence, swimming behaviour and mortality in a 3-dimensional
stochastic model. Marine Ecology Progress Series 311, 47–66.
Gunther, E.R. 1936. Variations in behaviour of the Perú Coastal Current: with an historical introduction. The
Geographical Journal 88, 37–61.
318
Guppy, H.B. 1906. Observations of a Naturalist in the Pacific between 1896 and 1899. New York: Macmillan.
Gutierrez, A., Correa, T., Muñoz, V., Santibañez, A., Marcos, R., Cáceres, C. & Buschmann, A.H. 2006.
Farming of the giant kelp Macrocystis pyrifera in southern Chile for development of novel food
products. Journal of Applied Phycology 18, 259–267.
Gutiérrez, D. 2000. Bioperturbación y macrofauna en fondos sublitorales de un área de surgencias frente a
Chile Central (36°30′S): variación espacial y temporal en el período 1997–1999. Ph.D. thesis,
Universidad de Concepción, Concepción, Chile.
Gutiérrez, D., Gallardo, V.A., Mayor, S., Neira, C., Vásquez, C., Sellanes, J., Rivas, M., Soto, A., Carrasco,
F. & Baltazar, M. 2000. Effects of dissolved oxygen and fresh organic matter on the bioturbation
potential of macrofauna in sublittoral sediments off Central Chile during the 1997/1998 El Niño.
Halpin, P.M., Strub, P.T., Peterson, W.T. & Baumgartner, T.R. 2004. An overview of interactions among
oceanography, marine ecosystems, climatic and human disruptions along the eastern margins of the
Pacific Ocean. Revista Chilena de Historia Natural 77, 371–409.
Hannach, G. & Santelices, B. 1985. Ecological differences between the isomorphic reproductive phases of
two species of Iridaea (Rodophyta, Gigartinales). Marine Ecology Progress Series 22, 219–303.
Harrold, C., Light, K. & Lisin, S. 1998. Organic enrichment of submarine-canyon and continental-shelf benthic
communities by macroalgal drift imported from nearshore kelp forests. Limnology and Oceanography
43, 669–678.
Harrold, C. & Pearse, J.S. 1987. The ecological role of echinoderms in kelps forest. In Echinoderm Studies,
M. Jangoux & J.M. Lawrence (eds). Rotterdam: Balkema, 137–233.
Hartmann-Schröder, G. & Hartmann, G. 1962. Zur Kenntnis des Eulitorals der chilenischen Pazifikküste und
der argentinischen Küste Südpatagoniens unter besonderer Berücksichtigung der Polychaeten und
Ostracoden. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, Ergänzungs-
band zu Band 60, 5–56.
Harzhauser, M., Piller, W.E. & Steininger, F.F. 2002. Circum-Mediterranean Oligo-Miocene biogeographic evolu-
tion — the gastropod’s point of view. Palaeogeography, Palaeoclimatology, Palaeoecology 183, 103–133.
Haury, L.R., McGowan, J.A. & Wiebe, P.H. 1978. Patterns and processes in the time-space scales of plankton
distributions. In Spatial Pattern in Plankton Communities, J.H. Steele (ed.). New York: Plenum Press,
277–327.
Hebbeln, D., Marchant, M., Freudenthal, T. & Wefer, G. 2000b. Surface sediment distribution along the Chilean
continental slope related to upwelling and productivity. Marine Geology 164, 119–137.
Hebbeln, D., Marchant, M. & Wefer, G. 2000a. Seasonal variations of the particle flux in the Peru-Chile
current at 30°S under ‘normal’ and El Niño conditions. Deep-Sea Research II 47, 2101–2128.
Heinrich, A.K. 1973. Horizontal distribution of copepods in the Perú Current region. Oceanology 13, 97–103.
Helly, J.J. & Levin, L.A. 2004. Global distribution of naturally occurring marine hypoxia on continental
margins. Deep-Sea Research I 51, 1159–1168.
Hennicke, J.C. & Culik, B.M. 2005. Foraging performance and reproductive success of Humboldt penguins
in relation to prey availability. Marine Ecology Progress Series 296, 173–181.
Hernández, C.E., Moreno, R.A. & Rozbaczylo, N. 2005. Biogeographical patterns and Rapoport’s rule in
southeastern Pacific benthic polychaetes of the Chilean coast. Ecography 28, 363–373.
Hernández, C.E., Muñoz, G. & Rozbaczylo, N. 2001. Poliquetos asociados con Austromegabalanus psittacus
(Molina, 1782) (Crustacea: Cirripedia) en Península Gualpén, Chile central: biodiversidad y efecto
del tamaño del sustrato biológico. Revista de Biología Marina y Oceanografía 36, 99–108.
Hernández, E.H. & Castro, L.R. 2000. Larval growth of the anchoveta Engraulis ringens during the winter
spawning season off central Chile. Fishery Bulletin 98, 704–710.
Hernández-Miranda, E., Palma, A.T. & Ojeda, F.P. 2003. Larval fish assemblages in nearshore coastal waters
off central Chile: temporal and spatial patterns. Estuarine Coastal and Shelf Science 56, 1075–1092.
Herrera, L. & Escribano, R. 2006. Factors structuring the phytoplankton community in the upwelling site off
El Loa River in northern Chile. Journal of Marine Systems 61, 13–38.
Herzka, S.Z., Holt, S.A. & Holt, G.J. 2002. Characterization of settlement patterns of red drum Sciaenops
ocellatus larvae to estuarine nursery habitat: a stable isotope approach. Marine Ecology Progress
Series 226, 143–156.
319
MARTIN THIEL ET AL.
Hidalgo, P. & Escribano, R. 2001. Succession of pelagic copepod species in coastal waters off northern Chile:
the influence of the 1997–998 El Niño. Hydrobiologia 453, 153–160.
Hidalgo, P. & Escribano, R. The coupling of life cycles of the copepods Calanus chilensis and Centropages
brachiatus to upwelling variability in the Central-South upwelling region off Chile (submitted).
Progress in Oceanography, Special Issue “The Structure and Functioning of the Coastal Upwelling
System off Central/south Chile.” R. Escribano & W. Schneider (eds).
Hidalgo, P., Escribano, R. & Morales, C.E. 2005a. Ontogenetic vertical distribution and diel migration of the
copepod Eucalanus inermis in the oxygen minimum zone off northern Chile (20–21°S). Journal of
Plankton Research 27, 519–529.
Hidalgo, P., Escribano, R. & Morales, C.E. 2005b. Annual life cycle of the copepod Eucalanus inermis at a
coastal upwelling site off Mejillones (23°S), northern Chile. Marine Biology 146, 995–1003.
Hilborn, R. & Walters, C.J. 1992. Quantitative Fisheries Stock Assessment. Choice, Dynamics and Uncertainty.
New York: Chapman and Hall.
Hill, A.E., Hickey, B.M., Shillington, F.A., Strub, P.T., Brink, K.H., Barton, E.D. & Thomas, A.C. 1998.
Eastern ocean boundaries. In The Sea, A.R. Robinson & K.H. Brink (eds). New York: John Wiley
and Sons, 29–68.
Hinojosa, I., Boltaña, S., Lancellotti, D., Macaya, E., Ugalde, P., Valdivia, N., Vásquez, N., Newman, W.A.
& Thiel, M. 2006. Geographic distribution and description of four pelagic barnacles along the south
east Pacific coast of Chile — a zoogeographical approximation. Revista Chilena de Historia Natural
79, 13–27.
Hinojosa, L.F. & Villagrán, C. 1997. Historia de los bosques templados del sur de Sudamérica, I: antecedentes
paleobotánicos, geológicos y climáticos del Terciario del cono sur de América. Revista Chilena de
Hoffmann, A.J. 1987. The arrival of seaweed propagules at the shore: a review. Botanica Marina 30, 151–165.
Hoffmann, A.J., Avila, M. & Santelices, B. 1984. Interaction of nitrate and phosphate on the development of
microscopic stages of Lessonia nigrescens Bory (Phaeophyta). Journal of Experimental Marine
Hoffmann, A.J. & Santelices, B. 1982. Effects of light intensity and nutrients on gametophytes and gameto-
genesis of Lessonia nigrescens Bory (Phaeophyta). Journal of Experimental Marine Biology and
Ecology 60, 77–89.
Hoffmann, A.J. & Santelices, B. 1991. Banks of algal microscopic forms — hypotheses on their functioning
and comparisons with seed banks. Marine Ecology Progress Series 79, 185–194.
Hoffmann, A. & Santelices, B. 1997. Flora Marina de Chile Central. Santiago, Chile: Ediciones Universidad
Católica de Chile.
Hoffmann, A.J. & Ugarte, R. 1985. The arrival of propagules of marine macroalgae in the intertidal zone.
Hofmann, G.E. & Somero, G.N. 1995. Evidence for protein damage at environmental temperatures, seasonal
changes in levels of ubiquitin conjugates and Hsp70 in the intertidal mussel Mytilus trossulus. Journal
of Experimental Biology 198, 1509–1518.
Hormazábal, S., Shaffer, G., Letelier, J. & Ulloa, O. 2001. Local and remote forcing of sea surface temperature
in the coastal upwelling system off Chile. Journal of Geophysical Research-Oceans 106, 16,657–16,671.
Hormazábal, S., Shaffer, G. & Leth, O. 2004. Coastal transition zone off Chile. Journal of Geophysical
Research-Oceans 109, C01021 DOI:10.1029/2003JC001956.
Hückstädt, L.A. & Antezana, T. 2003. Behaviour of the southern sea lion (Otaria flavescens) and consumption
of the catch during purse-seining for jack mackerel (Trachurus symmetricus) off central Chile. ICES
Journal of Marine Science 60, 1003–1011.
Hückstädt, L.A. & Antezana, T. 2004. Behaviour of southern sea lions in presence of killer whales during
fishing operations in central Chile. Scientia Marina 68, 295–298.
Hückstädt, L.A. & Krautz, M.C. 2003. Interaction between southern sea lions Otaria flavescens and jack
mackerel Trachurus symmetricus commercial fishery off central Chile: a geostatistical approach.
Humboldt, A. 1846. Cosmos: A Sketch of a Physical Description of the Universe. London: Green and
Longmans.
320
Huovinen, P., Gómez, I., Figueroa, F.L., Ulloa, N., Morales, V. & Lovengreen, C. 2004. Ultraviolet absorbing
mycosporine like amino acids in red macroalgae from Chile. Botanica Marina 47, 21–29.
Huovinen, P., Gómez, I. & Lovengreen, C. 2006. A five year study of solar ultraviolet radiation in southern
Chile (39°S): potential impact on physiology of coastal marine algae? Photochemistry and Photobi-
ology 82, 515–522.
Hutchins, D.A., Hare, C.E., Weaver, R.S., Zhang, Y., Firme, G.F., DiTullio, G.R., Alm, M.B., Riseman, S.F.,
Maucher, J.M., Geesey, M.E., Trick, C.G., Smith, G.J., Rue, E.L., Conn, J. & Bruland, K.W. 2002.
Phytoplankton iron limitation in the Humboldt Current and Perú Upwelling. Limnology and Ocean-
ography 47, 997–1011.
Huyer, A., Smith, R.L. & Paluszkiewicz, T. 1987. Coastal upwelling off Perú during normal and El Niño
times, 1981–1984. Journal of Geophysical Research-Oceans 92, 14,297–14,307.
Ianora, A., Miralto, A., Poulet, S.A., Carotenuto, Y., Buttino, I., Romano, G., Casotti, R., Pohnert, G., Wichard,
T., Colucci-D’Amato, L., Terrazzano, G. & Smetacek, V. 2004. Aldehyde suppression of copepod
recruitment in blooms of a ubiquitous planktonic diatom. Nature 429, 403–407.
Ibáñez, C.M., González, C. & Cubillos, L. 2004. Dieta del pez espada Xiphias gladius Linnaeus, 1758, en
aguas oceánicas de Chile central en invierno de 2003. Investigaciones Marinas, Valparaíso 32,
113–120.
IFOP. 1977. Evaluación del recurso algas pardas en la I Región. Informe Final SERPLAC I Región. Iquique,
Chile: IFOP.
Illanes, J.E., Akaboshi, S. & Uribe, E. 1985. Temperature effects on reproduction of the scallop Chlamys
(Argopecten) purpuratus in Bahía Tongoy, during the 1982–1983 El Niño phenomenon. Investiga-
ciones Pesqueras (Chile) 32, 167–173.
IMARPE. 2006. Aves guaneras: situación actual y perspectivas. Available HTTP: http://www.imarpe.gob.pe/
aves_marinas.html. Accessed 21 February 2007.
Iriarte, J.L. & Bernal, P. 1990. Vertical distribution of diatoms and thecate dinoflagellates in the Gulf of Arauco:
species composition, relative abundance, and the chlorophyll maximum layer. Scientia Marina 54,
389–399.
Iriarte, J.L. & González, H.E. 2004. Phytoplankton size structure during and after the 1997/1998 El Niño in
a coastal upwelling area of the northern Humboldt Current System. Marine Ecology Progress Series
269, 83–90.
Iriarte, J.L., Pizarro, G., Troncoso, V.A. & Sobarzo, M. 2000. Primary production and biomass of size-
fractionated phytoplankton off Antofagasta, Chile (23–24°S) during pre-El Niño and El Niño 1997.
Journal of Marine Systems 26, 37–51.
IUCN. 1994. Guidelines for Protected Area Management Categories. Gland, Switzerland: IUCN.
IUCN. 2006. IUCN Red List of Threatened Species. Available HTTP: http://www.iucnredlist.org. Accessed
21 February 2007.
Jahncke, J., Checkley, D.M. & Hunt, G.L. 2004. Trends in carbon flux to seabirds in the Peruvian upwelling
system: effects of wind and fisheries on population regulation. Fisheries Oceanography 13, 208–223.
Jara, C.G. 1997. Antecedentes sobre el desarrollo de la carcinología en Chile. Investigaciones Marinas,
Valparaíso 25, 245–254.
Jara, F. & Moreno, C.A. 1983. Calendario de reclutamiento de organismos epibénticos móviles de la zona
mesomareal de Mehuín, Chile. Medio Ambiente 6, 72–79.
Jara, H.F. & Moreno, C.A. 1984. Hervibory and structure in a mid-littoral rocky community: a case in southern
Chile. Ecology 65, 28–38.
Jaramillo, E. 1982. Taxonomy, natural history and zoogeography of sand beach isopods from the coast of
southern Chile. Studies on Neotropical Fauna and Environment 17, 175–194.
Jaramillo, E. 1987. Sandy beach macroinfauna from the Chilean coast: zonation patterns and zoogeography.
Vie et Milieu 37, 165–174.
Jaramillo, E. 1994. Patterns of species richness in sandy beaches of South America. South African Journal of
Zoology-Suid-Afrikaanse Tydskrif Vir Dierkunde 29, 227–234.
Jaramillo, E., Avellanal, M.H., González, M. & Kennedy, F. 2000. Locomotor activity of Phalerisida maculata
Kulzer (Coleoptera, Tenebrionidae) on Chilean sandy beaches. Revista Chilena de Historia Natural
73, 67–77.
321
MARTIN THIEL ET AL.
Jaramillo, E., Carrasco, F., Quijón, P., Pino, M. & Contreras, H. 1998. Distribution and community structure
of the benthic macroinfauna at the coast of northern Chile. Revista Chilena de Historia Natural 71,
459–478.
Jaramillo, E., Contreras, H., Duarte, C. & Avellanal, M.H. 2003. Locomotor activity and zonation of upper shore
arthropods in a sandy beach of north central Chile. Estuarine Coastal and Shelf Science 58, 177–197.
Jaramillo, E., Contreras, H., Duarte, C. & Quijón, P. 2001. Relationships between community structure of the
intertidal macroinfauna and sandy beach characteristics along the Chilean coast. Marine Ecology
Pubblicazioni Della Stazione Zoologica Di Napoli I 22, 323–342.
Jaramillo, E., Stotz, W., Bertrán, C., Navarro, J., Román, C. & Varela, C. 1980. Actividad locomotriz de
Orchestoidea tuberculata (Amphipoda, Talitridae) sobre la superficie de una playa arenosa del Sur
de Chile (Mehuín, Provincia de Valdivia). Studies on Neotropical Fauna and Environment 15, 9–33.
Jerardino, A., Castilla, J.C., Ramírez, J.M. & Hermosilla, N. 1992. Early coastal substince patterns in Central
Chile: a systematic study of the marine invertebrate fauna from the site of Curahumilla 1. Latin
American Antiquity 3, 43–62.
Jesse, S. & Stotz, W. 2002. Spatio-temporal distribution patterns of the crab assemblage in the shallow subtidal
of the north Chilean Pacific coast. Crustaceana 75, 1161–1200.
Jiménez, M. 2005. Bioecología de Arctocephalus australis en la península de Mejillones, II Región, Chile.
Honors thesis, Universidad de Antofagasta, Antofagasta, Chile.
Johnson, D.D. 1995. The United States in the Pacific: Private Interests and Public Policies, 1784–1899.
Westport, CT: Praeger Publishers.
Johnson, D.R., Fonseca, T. & Sievers, H. 1980. Upwelling in the Humboldt coastal current near Valparaíso,
Chile. Journal of Marine Research 38, 1–36.
Jones, C.G., Lawton, J.H. & Shachak, M. 1994. Organisms as ecosystem engineers. Oikos 69, 373–386.
Kain, J. 1989. The seasons in the subtidal. British Phycology J 24, 203–215.
Kamykowski, D. & Zentara, S.J. 1990. Hypoxia in the world ocean as recorded in the historical data set.
Deep-Sea Research A 37, 1861–1874.
Kaplan, D.M. 2006. Alongshore advection and marine reserves: consequences for modeling and management.
Kaplan, D.M., Largier, J.L., Navarrete, S., Guiñez, R. & Castilla, J.C. 2003. Large diurnal temperature
fluctuations in the nearshore water column. Estuarine Coastal and Shelf Science 57, 385–398.
Kaufman, L., Jackson, J.B., Sala, E., Chisolm, P., Gomez, E.D., Peterson, C., Salm, R.V. & Llewellyn, G.
2004. Restoring and maintaining marine ecosystem function. In Defying Ocean’s End: An Agenda
for Action, L.K. Glover & S.A. Earle (eds). Washington, DC: Island Press, 165–181.
Keefer, D.K., deFrance, S.D., Moseley, M.E., Richardson, J.B., Satterlee, D.R. & Day-Lewis, A. 1998. Early
maritime economy and El Niño events at Quebrada Tacahuay, Perú. Science 281, 1833–1835.
Kelaher, B.P. & Castilla, J.C. 2005. Habitat characteristics influence macrofaunal communities in coralline
turf more than mesoscale coastal upwelling on the coast of northern Chile. Estuarine Coastal and
Keough, M.J. & Downes, B.J. 1982. Recruitment of marine invertebrates: the role of active larval choices and
early mortality. Oecologia 54, 248–352.
Kerr, R.A. 1999. El Niño grew strong as cultures were born. Science 283, 467–468.
Kim, S.L. 1992. The role of drift kelp in the population ecology of a Diopatra ornata Moore (Polychaeta,
Onuphidae) ecotone. Journal of Experimental Marine Biology and Ecology 156, 253–272.
Kinlan, B.P. & Gaines, S.D. 2003. Propagule dispersal in marine and terrestrial environments: a community
perspective. Ecology 84, 2007–2020.
Klinger, T., Padilla, D.K. & Britton-Simmons, K. 2006. Two invaders achieve higher densities in reserves.
Aquatic Conservation-Marine and Freshwater Ecosystems 16, 301–311.
Knox, G.A. 1960. Littoral ecology and biography of the southern oceans. Proceedings of the Royal Society
Series B Biological Sciences 152, 577–624.
Kortum, G. 2002. Germania in Pacífico: Humboldt, Chamisso and other early German contributions to Pacific
research, 1741–1876. In Oceanographic History: the Pacific and Beyond, K.R. Benson & P.F. Rehbock
(eds). Seattle: University of Washington Press, 107–117.
322
Kroencke, I., Dippner, J.W., Heyen, H. & Zeiss, B. 1998. Long-term changes in macrofaunal communities
off Norderney (East Frisia, Germany) in relation to climate variability. Marine Ecology Progress
Series 167, 25–36.
Kuypers, M.M., Lavik, G., Woebken, D., Schmid, M., Fuchs, B.M., Amann, R., Jorgensen, B.B. & Jetten,
M.S.M. 2005. Massive nitrogen loss from the Benguela upwelling system through anaerobic ammo-
nium oxidation. Proceedings of the National Academy of Sciences USA 102, 6478–6483.
Ladah, L.B., Zertuche-González, J.A. & Hernández-Carmona, G. 1999. Giant kelp (Macrocystis pyrifera,
Phaeophyceae) recruitment near its southern limit in Baja California after mass disappearance during
ENSO 1997–1998. Journal of Phycology 35, 1106–1112.
Lagos, N.A., Barria, I. & Paolini, P. 2002. Upwelling ecosystem of northern Chile: integrating benthic ecology
and coastal oceanography through remote sensing. In The Oceanography and Ecology of the Nearshore
and Bays in Chile, J.C. Castilla & J. Largier (eds). Santiago, Chile: Ediciones Universidad Católica
de Chile, 117–141.
Lagos, N.A., Navarrete, S.A., Véliz, F., Masuero, A. & Castilla, J.C. 2005. Meso-scale spatial variation in
settlement and recruitment of intertidal barnacles along the coast of central Chile. Marine Ecology
Lagos, N.A., Tapia, F., Navarrete, S.A. & Castilla, J.C. In review. Synchrony in recruitment of benthic
invertebrates in central Chile. Oikos.
Lancellotti, D.A. 2002. Midiendo cambios en comunidades macrobentónicas marinas. M.Sc. thesis, Univer-
sidad Católica del Norte, Coquimbo, Chile.
Lancellotti, D.A. & Stotz, W.B. 2004. Effects of shoreline discharge of iron mine tailings on a marine soft-
bottom community in northern Chile. Marine Pollution Bulletin 48, 303–312.
Lancellotti, D.A. & Vásquez, J.A. 1999. Biogeographical patterns of benthic macroinvertebrates in the south-
eastern Pacific littoral. Journal of Biogeography 26, 1001–1006.
Lancellotti, D.A. & Vásquez, J.A. 2000. Zoogeography of benthic macroinvertebrates of the Chilean coast:
contribution for marine conservation. Revista Chilena de Historia Natural 73, 99–129.
Landaeta, M.F. & Castro, L.R. 2002. Spring spawning and early nursery zone of the mesopelagic fish
Maurolicus parvipinnis at the coastal upwelling zone off Talcahuano, central Chile. Marine Ecology
Landaeta, M.F. & Castro, L.R. 2006. Spawning and larval survival of the Chilean hake Merluccius gayi under
later summer conditions in the Gulf of Arauco, central Chile. Fisheries Research 77, 115–121.
Landaeta, M.F., Herrera, G.A., Pedraza, M., Bustos, C. & Castro, L.R. 2006. Reproductive tactics and larval
development of bigeye flounder, Hippoglossina macrops, off central Chile. Journal of the Marine
Biological Association of the United Kingdom 86, 1253–1264.
Lardies, M.A. & Castilla, J.C. 2001. Latitudinal variation in the reproductive biology of the commensal crab
Pinnaxodes chilensis (Decapoda: Pinnotheridae) along the Chilean coast. Marine Biology 139, 1125–1133.
Largier, J.L. 2002. Linking oceanography and nearshore ecology: perspectives and challenges. In The Ocean-
ography and Ecology of the Nearshore and Bays in Chile, J.C. Castilla & J.L. Largier (eds). Santiago,
Chile: Ediciones Universidad Católica de Chile, 117–141.
Largier, J.L. 2003. Considerations in estimating larval dispersal distances from oceanographic data. Ecological
Applications 13, S71–S89.
Leber, K.M. 1982. Bivalves (Tellinacea: Donacidae) on a North Carolina beach: contrasting population size
structures and tidal migrations. Marine Ecology Progress Series 7, 297–301.
Lee, C.E. & Strathmann, R.R. 1998. Scaling of gelatinous clutches: effects of siblings’ competition for oxygen
on clutch size and parental investment per offspring. American Naturalist 151, 293–310.
Le Fèvre, J. & Bourget, E. 1992. Hydrodynamics and behavior — transport processes in marine invertebrate
larvae. Trends in Ecology and Evolution 7, 288–289.
Leibold, M.A., Holyoak, M., Mouquet, N., Amarasekare, P., Chase, J.M., Hoopes, M.F., Holt, R.D., Shurin,
J.B., Law, R., Tilman, D., Loreau, M. & González, A. 2004. The metacommunity concept: a framework
for multi-scale community ecology. Ecology Letters 7, 601–613.
Lemmens, J., Clapin, G., Lavery, P. & Cary, J. 1996. Filtering capacity of seagrass meadows and other habitats
of Cockburn Sound, western Australia. Marine Ecology Progress Series 143, 187–200.
323
MARTIN THIEL ET AL.
León, R.I. & Stotz, W. 2004. Diet and prey selection dynamics of Cancer polyodon in three different habitat types
in Tongoy Bay, Chile. Journal of the Marine Biological Association of the United Kingdom 84, 751–756.
Leslie, H., Breck, E., Chan, F., Lubchenco, J. & Menge, B.A. 2005. Barnacle reproductive hotspots linked to
nearshore ocean conditions. Proceedings of the National Academy USA 102, 10,534–10,539.
Leslie, H., Ruckelshaus, M., Ball, I.R., Andelman, S. & Possingham, H.P. 2003. Using siting algorithms in
the design of marine reserve networks. Ecological Applications 13, S185–S198.
Letelier, J., Pizarro, O., Núñez, S. & Arcos, D. 2004. Spatial and temporal variability of thermal fronts off
central Chile (33–40°S). Gayana (Concepción) 38, 358–362.
Letelier, S., Vega, M.A., Ramos, A.M. & Carreno, E. 2003. Molluscs from Chile in the Chilean national
museum of natural history database. Revista de Biología Tropical 51(Suppl. 3), 33–137.
Leth, O., Shaffer, G. & Ulloa, O. 2004. Hydrography of the eastern South Pacific Ocean: results from the
Sonne 102 cruise, May–June 1995. Deep-Sea Research II 51, 2349–2369.
Levin, L.A. 2002. Deep-ocean life where oxygen is scarce. American Scientist 90, 436–444.
Levin, L.A. 2003. Oxygen minimum zone benthos: adaptation and community response to hypoxia. Ocean-
ography and Marine Biology: An Annual Review 41, 1–45.
Levin, L.A. 2006. Recent progress in understanding larval dispersal: new directions and digressions. Integrative
and Comparative Biology 46, 282–297.
Levin, L.A., Etter, R.J., Rex, M.A., Gooday, A.J., Smith, C.R., Pineda, J., Stuart, C.T., Hessler, R.R. & Pawson,
D. 2001. Environmental influences on regional deep-sea species diversity. Annual Review of Ecology
and Systematics 32, 51–93.
Levin, L.A. & Gage, J.D. 1998. Relationships between oxygen, organic matter and the diversity of bathyal
macrofauna. Deep-Sea Research II 45, 129–163.
Levin, L.A., Gage, J.D., Martin, C. & Lamont, P.A. 2000. Macrobenthic community structure within and
beneath the oxygen minimum zone, NW Arabian Sea. Deep-Sea Research II 47, 189–226.
Levin, L.A., Huggett, C.L. & Wishner, K.F. 1991. Control of deep-sea benthic community structure by oxygen
and organic-matter gradients in the eastern Pacific Ocean. Journal of Marine Research 49, 763–800.
Livingstone, D.R. 1983. Invertebrate and vertebrate pathway of anaerobic metabolism: evolutionary consid-
erations. Journal of the Geological Society 140, 27–37.
Llagostera, A. 1979. 9700 years of maritime subsistence on the Pacific: an analysis by means of bioindicators
in the north of Chile. American Antiquity 44, 309–323.
Llanos, A., Herrera, G. & Bernal, P. 1996. Análisis del tamaño de presas en la dieta de las larvas de cuatro
Clupeiformes en un área costera de Chile central. Scientia Marina 60, 435–442.
Llanos-Rivera, A. & Castro, L.R. 2004. Latitudinal and seasonal egg-size variation of the anchoveta (Engraulis
ringens) off the Chilean coast. Fishery Bulletin 102, 207–212.
Llanos-Rivera, A. & Castro, L.R. 2006. Inter-population differences in temperature effects on Engraulis ringens
yolk-sac larvae. Marine Ecology Progress Series 312, 245–253.
Llanos-Rivera, A., Herrera, G. & Bernal, P. 2004. Food size selectivity and diet overlap in larvae of Clupeiform
species from central Chile. Cahiers De Biologie Marine 45, 1–8.
Lleellish, J., Fernández, E. & Hooker, Y. 2001. Disturbancia del bosque submareal de Macrocystis pyrifera
durante El Niño 1997–1998 en la Bahía de Pucusana. In Sustentabilidad de la Biodiversidad. Un
Problema Actual: Bases Científico Técnicas, Teorizaciones y Proyecciones, K. Alveal & T. Antezana
(eds). Concepción, Chile: Universidad de Concepción, 331–350.
Lockwood, D.R., Hastings, A. & Botsford, L.W. 2002. The effects of dispersal patterns on marine reserves:
does the tail wag the dog? Theoretical Population Biology 61, 297–309.
Loeb, V.J. & Rojas, O. 1988. Interannual variation of ichthyoplankton composition and abundance relations
off northern Chile, 1964–1983. Fishery Bulletin 86, 1–24.
López, C.A. & Stotz, W.B. 1997. Description of the fauna associated to Corallina officinalis L. in the intertidal
of the rocky shore of ‘Palo Colorado’ (Los Vilos, IV Región, Chile). Revista de Biología Marina y
Oceanografía 32, 17–35.
Lothrop, S.K. 1932. Aboriginal navigation off the west coast of south America. The Journal of the Royal
Anthropological Institute of Great Britain and Ireland 62, 229–256.
Ludynia, K., Garthe, S. & Luna-Jorquera, G. 2005. Seasonal and regional variation in the diet of the Kelp
gull in northern Chile. Waterbirds 28, 359–365.
324
Luna-Jorquera, G. & Culik, B.M. 1999. Diving behaviour of Humboldt penguins Spheniscus humboldti in
northern Chile. Marine Ornithology 27, 67–76.
Luna-Jorquera, G., Simeone, A. & Aguilar, R. 2003. Ecofisiología de animales endotermos en un desierto
cálido y un mar frío: el caso de las aves marinas de la corriente de Humboldt. In Fisiología Ecológica
y Evolutiva. Teoría y Casos de Estudios en Animales, F. Bozinovic (ed.). Santiago, Chile: Ediciones
Universidad Católica de Chile, 297–316.
Lüning, K. 1990. Seaweeds. Their Enviroment, Biogeography, and Ecophysiology. New York: Wiley-Interscience.
Lüning, K. & tom Dieck, I. 1989. Environmental triggers in algal seasonality. Botanica Marina 32, 389–397.
Macaya, E.C., Boltaña, S., Hinojosa, I.A., Macchiavello, J.E., Valdivia, N.A., Vásquez, N.R., Buschmann,
A.H., Vásquez, J.A., Vega, J.M.A. & Thiel, M. 2005. Presence of sporophylls in floating kelp rafts
of Macrocystis spp. (Phaeophyceae) along the Chilean Pacific coast. Journal of Phycology 41, 913–922.
Macchiavello, J., Bulboa, C.R. & Edding, M. 2003. Vegetative propagation and spore-based recruitment in
the carragenophyte Chondracanthus chamissoi (Gigartinales, Rhodophyta) in northern Chile. Phyco-
logical Research 51, 45–50.
MacDonald, B.A. & Thompson, R.J. 1985. Influence of temperature and food availability on the ecological
energetics of the giant scallops Placopecten magellanicus. II. Reproductive output and total produc-
tion. Marine Ecology Progress Series 116, 295–303.
MacDonald, B.A. & Thompson, R.J. 1988. Intraspecific variation in growth and reproduction in latitudinally
differentiated populations of the giant scallops Placopecten magellanicus. Biological Bulletin 175,
361–371.
Madden, R.A. & Julian, P.R. 1971. Detection of a 40–50 day oscillation in the zonal wind in the tropical
Pacific. Journal of Atmospheric Sciences 28, 702–708.
Madronich, S., McKenzie, R.L., Caldwell, M. & Bjorn, L.O. 1995. Changes in ultraviolet radiation reaching
the Earth’s surface. Ambio 24, 143–152.
Madsen, F.J. 1956. Asteroidea, with a survey of the Asteroidea of the Chilean shelf. Lunds Universitets Arsskrift
N.F. 52, 1–53.
Majluf, P., Babcock, E.A., Riveros, J.C., Schreiber, M.A. & Alderete, W. 2002. Catch and bycatch of sea birds
and marine mammals in the small-scale fishery of Punta San Juan, Peru. Conservation Biology 16,
1333–1343.
Maldonado, A. & Villagrán, C. 2002. Paleoenvironmental changes in the semiarid coast of Chile (~32°S)
during the last 6200 cal years inferred from a swamp-forest pollen record. Quaternary Research 58,
130–138.
Mann, G. 1954. Vida de los Peces en Aguas Chilenas. Santiago, Chile: Instituto de Investigaciones Veterinarias,
Universidad de Chile.
Manríquez, P.H. & Castilla, J.C. 2007. Roles of larval behaviour and microhabitat traits in determining spatial
aggregations in the ascidian Pyura chilensis. Marine Ecology Progress Series 332, 155–165.
Manríquez, P.H., Navarrete, S.A., Rosson, A. & Castilla, J.C. 2004. Settlement of the gastropod Concholepas
concholepas on shells of conspecific adults. Journal of the Marine Biological Association of the United
Kingdom 84, 651–658.
Marchant, M., Hebbeln, D., Giglio, S., Coloma, C. & González, H.E. 2004. Seasonal and interannual variability
in the flux of planktic foraminifera in the Humboldt Current System off central Chile (30°S). Deep-
Sea Research II 51, 2441–2455.
Marchesiello, P., McWilliams, J.C. & Shchepetkin, A. 2001. Open boundary condition for long-term integration
of regional oceanic models. Ocean Modelling 3, 1–21.
Marchesiello, P., McWilliams, J.C. & Shchepetkin, A. 2003. Equilibrium structure and dynamics of the
California Current System. Journal of Physical Oceanography 33, 753–783.
Marcus, N.H. & Boero, F. 1998. Minireview: the importance of benthic-pelagic coupling and the forgotten
role of life cycles in coastal aquatic systems. Limnology and Oceanography 43, 763–768.
Marín, V.H. 1997. A simple-biology, stage-structured population model of the spring dynamics of Calanus
chilensis at Mejillones del Sur Bay, Chile. Ecological Modelling 105, 65–82.
Marín, V.H. 2000. Large-scale patterns and processes in marine ecosystems: hypotheses and methods. In
Antarctic Ecosystems: Models for Wider Ecological Understanding, W. Davidson et al. (eds). Christ-
church, New Zealand: The Caxton Press, 197–202.
325
MARTIN THIEL ET AL.
Marín, V.H. & Delgado, L.E. 2007. Lagrangian observations of surface coastal flows north of 30°S in the
Humboldt Current System. Continental Shelf Research 27, 731–743.
Marín, V.H., Delgado, L.E. & Escribano, R. 2003b. Upwelling shadows at Mejillones Bay (northern Chilean
coast): a remote sensing in situ analysis. Investigaciones Marinas, Valparaíso 31, 47–55.
Marín, V.H., Delgado, L.E. & Luna-Jorquera, G. 2003a. S-Chlorophyll squirts at 30°S off the Chilean coast
(eastern South Pacific): feature-tracking analysis. Journal of Geophysical Research-Oceans 108, C12,
3378, DOI:10.1029/2003JC001935.
Marín, V.H., Escribano, R., Delgado, L.E., Olivares, G. & Hidalgo, P. 2001. Nearshore circulation in a coastal
upwelling site off the northern Humboldt Current System. Continental Shelf Research 21, 1317–1329.
Marín, V.H., Espinoza, S. & Fleminger, A. 1994. Morphometric study of Calanus chilensis males along the
Chilean coast. Hydrobiologia 293, 75–80.
Marín, V.H. & Moreno, C.A. 2002. Wind driven circulation and larval dispersal: a review of its consequences
in coastal benthic recruitment. In The Oceanography and Ecology of the Nearshore and Bays in Chile,
J.C. Castilla & J.L. Largier (eds). Santiago, Chile: Ediciones Universidad Católica de Chile, 47–63.
Marín, V.H. & Olivares, G.R. 1999. Seasonality of primary productivity in Mejillones del Sur Bay (Chile): a
process-functional approach. Revista Chilena de Historia Natural 72, 629–641.
Marín, V.H., Rodríguez, L., Vallejo, L., Fuenteseca, J. & Oyarce, E. 1993. Effects of coastal upwelling on the
spring primary productivity of Mejillones del sur Bay (Antofagasta, Chile). Revista Chilena de Historia
Natural 66, 479–491.
Marincovich, L., Jr. 1973. Intertidal mollusks of Iquique, Chile. Natural History Museum of Los Angeles
County Science Bulletin 16, 1–49.
Marquet, P.A., Fernández, M., Navarrete, S. & Valdovinos, C. 2004. Diversity emerging: toward a deconstruc-
tion of biodiversity patterns. In Frontiers of Biogeography, M.V. Lomolino & L.R. Heaney (eds).
Sunderland, Massachusetts: Sinauer Associates, 191–209.
Martin, J.H., Fitzwater, S.E., Gordon, R.M., Hunter, C.N. & Tanner, S.J. 1993. Iron, primary production and
carbon nitrogen flux studies during the JGOFS North-Atlantic bloom experiment. Deep-Sea
Research II 40, 115–134.
Martin, J.H. & Gordon, R.M. 1988. Northeast Pacific iron distributions in relation to phytoplankton produc-
tivity. Deep-Sea Research A 35, 177–196.
Martínez, E.A. 1999. Latitudinal differences in thermal tolerance among microscopic sporophytes of the kelp
Lessonia nigrescens (Phaeophyta: Laminariales). Pacific Science 53, 74–81.
Martínez, E.A., Cardenas, L. & Pinto, R. 2003. Recovery and genetic diversity of the intertidal kelp Lessonia
nigrescens (Phaeophyceae) 20 years after El Niño 1982/83. Journal of Phycology 39, 504–508.
Martínez, E.A. & Santelices, B. 1998. Selective mortality on haploid and diploid microscopic stages of
Lessonia nigrescens Bory (Phaeophyta, Laminariales). Journal of Experimental Marine Biology and
Ecology 229, 219–239.
Martínez, G., Brokordt, K., Aguilera, C., Soto, V. & Guderley, H. 2000. Effect of diet and temperature upon
muscle metabolic capacities and biochemical composition of gonad and muscle in Argopecten pur-
puratus Lamarck 1819. Journal of Experimental Marine Biology and Ecology 247, 29–49.
Martínez, P. & Navarrete, S.A. 2002. Temporal and spatial variation in settlement of the gastropod Concholepas
concholepas in natural and artificial substrata. Journal of the Marine Biological Association of the
United Kingdom 82, 257–264.
Mascia, M. 2004. Social dimensions of marine reserves. In Marine Reserves: A Guide to Science, Design and
Use, J.A. Sobel & C.P. Dahlgren (eds). Washington, D.C.: Island Press, 164–186.
Mattern, T., Ellenberg, U. & Luna-Jorquera, G. 2002. A South American marine otter Lontra felina preys
upon chicks of the Peruvian diving petrel Pelecanoides garnotii. Marine Ornithology 30, 95–96.
McLachlan, A. & Jaramillo, E. 1995. Zonation on sandy beaches. Oceanography and Marine Biology: An
Annual Review 33, 305–335.
Medina, M. & Arancibia, H. 2002. Dinámica trófica del jurel (Trachurus symmetricus murphyi) en el norte
de Chile. Investigaciones Marinas, Valparaíso 30, 45–55.
Medina, M., Araya, M. & Vega, C. 2004. Alimentación y relaciones tróficas de peces costeros de la zona norte
de Chile. Investigaciones Marinas, Valparaíso 32, 33–47.
326
Meir, E., Andelman, S. & Possingham, H.P. 2004. Does conservation planning matter in a dynamic and
uncertain world? Ecology Letters 7, 615–622.
Meltzoff, S.K., Lichtensztajn, Y.G. & Stotz, W. 2002. Competing visions for marine tenure and co-management:
genesis of a marine management area system in Chile. Coastal Management 30, 85–99.
Méndez, C.A. 2002. Cazadores recolectores costeros y sus contextos de tarea: una división desde el asentam-
iento holocénico temprano de Punta Penitente (Lv 0.14), Los Vilos. Chungará (Arica) 34, 135–166.
Méndez, C.A. & Jackson, D.G. 2004. Ocupaciones humanas del holoceno tardío en Los Vilos (IV Región,
Chile): origen y características conductuales de la población local de cazadores recolectores de litoral.
Chungará (Arica) 36, 279–293.
Meneses, I. 1993. Foam as a dispersal agent in the rocky intertidal of central Chile. European Journal of
Meneses, I. & Santelices, B. 2000. Patterns and breaking points in the distribution of benthic algae along the
temperate Pacific coast of South America. Revista Chilena de Historia Natural 73, 615–623.
Menge, B.A. 1972. Foraging strategy of a starfish in relation to actual prey availability and environmental
predictability. Ecological Monographs 42, 25–50.
Menge, B.A., Berlow, E.L., Blanchette, C.A., Navarrete, S.A. & Yamada, S.B. 1994. The keystone species concept
— variation in interaction strength in a rocky intertidal habitat. Ecological Monographs 64, 249–286.
Menge, B.A., Lubchenco, J., Bracken, M.E.S., Chan, F., Foley, M.M., Freidenburg, T.L., Gaines, S.D., Hudson,
G., Krenz, C., Leslie, H., Menge, D.N.L., Russell, R. & Webster, M.S. 2003. Coastal oceanography
sets the pace of rocky intertidal community dynamics. Proceedings of the National Academy of
Sciences USA 100, 12,229–12,234.
Menzies, R.J. 1962. The zoogeography, ecology and systematics of the Chilean marine isopods. Lunds
Universitets Arsskrift NF 57, 1–162.
Mesías, J.M., Matano, R.P. & Strub, P.T. 2001. A numerical study of the upwelling circulation off central
Chile. Journal of Geophysical Research-Oceans 106, 19,611–19,623.
Mesías, J.M., Matano, R.P. & Strub, P.T. 2003. Dynamical analysis of the upwelling circulation off central
Chile. Journal of Geophysical Research-Oceans 108, C3, 3085 DOI:101029/2001JC 001135.
Micheli, F., Amarasekare, P., Bascompte, J. & Gerber, L.R. 2004. Including species interactions in the design
and evaluation of marine reserves: some insights from a predator-prey model. Bulletin of Marine
Science 74, 653–669.
Milessi, A., Sellanes, J., Gallardo, V. & Lange, C.B. 2005. Osseous skeletal material and fish scales in marine
sediments under the oxygen minimum zone off northern and central Chile. Estuarine, Coastal and
Miranda, C. 2001. La desaparición del banco de machas Mesodesma donacium (Lamarck, 1818) (Mollusca:
Bivalvia: Mesodesmatidae) en la Bahía de Coquimbo, IV Región, Chile: sus probables causas. Honors
Moloney, C.L., Jarre, A., Arancibia, H., Bozec, Y.M., Neira, S., Roux, J.P. & Shannon, L.J. 2005. Comparing
the Benguela and Humboldt marine upwelling ecosystems with indicators derived from inter-calibrated
models. ICES Journal of Marine Science 62, 493–502.
Montecino, V., Astoreca, R., Alarcón, G., Retamal, L. & Pizarro, G. 2004. Bio-optical characteristics and
primary productivity during upwelling and non-upwelling conditions in a highly productive coastal
ecosystem off central Chile (~36°S). Deep-Sea Research II 51, 2413–2426.
Montecino, V. & Pizarro, G. 1995. Phytoplankton acclimation and spectral penetration of UV irradiance off
the central Chilean coast. Marine Ecology Progress Series 121, 261–269.
Montecino, V., Pizarro, G. & Quiroz, D. 1996. Dinámica fitoplanctónica en el sistema de surgencia frente a
Coquimbo (30°S) a través de la relación funcional entre fotosíntesis e irradianza (P-I). Gayana
Oceanológica 4, 139–151.
Montecino, V. & Quiroz, D. 2000. Specific primary production and phytoplankton cell size structure in an
upwelling area off the coast of Chile (30°S). Aquatic Sciences 62, 364–380.
Montecino, V., Strub, P.T., Chavez, F.P., Thomas, A.C., Tarazona, J. & Baumgartner, T. 2005. Chapter 10
Bio-physical interactions off western South America (6,E). In The Sea, A.R. Robinson & K.H. Brink
(eds). Cambridge: Harvard University Press, 329–390.
327
MARTIN THIEL ET AL.
Montecinos, A., Purca, S. & Pizarro, O. 2003. Interannual-to-interdecadal sea surface temperature variability
along the western coast of South America. Geophysical Research Letters 30, 11, 1570, DOI:10.1029/
2003GL017345.
Moore, P.G. 2002. Mammals in intertidal and maritime ecosystems: interactions, impacts and implications.
Moraga, J., Valdebenito, E. & Rutllant, J. 2001. Condiciones oceanográficas durante la fase de relajación de
un evento de surgencia invernal frente a Punta Lengua de Vaca, Coquimbo. Investigaciones Marinas,
Valparaíso 29, 59–71.
Moragat, D., Avendaño, M., Peña, J., Le Pennect, M., Tanguyt, A. & Baron, J. 2001. Genetic and morphological
differentiation between two pectinid populations of Argopecten purpuratus from the northern Chilean
coast. Estudios Oceanológicos 20, 51–60.
Morales, C.E. 1999. Carbon and nitrogen fluxes in the oceans: the contribution by zooplankton migrants to
active transport in the North Atlantic during the Joint Global Ocean Flux Study. Journal of Plankton
Research 21, 1799–1808.
Morales, C.E., Blanco, J.L., Braun, M., Reyes, H. & Silva, N. 1996. Chlorophyll-a distribution and associated
oceanographic conditions in the upwelling region off northern Chile during the winter and spring
1993. Deep-Sea Research I 43, 267–289.
Morales, C.E., González, H.E., Hormazábal, S.E., Yuras, G., Letelier, J. & Castro, L.R. Accepted. The
distribution of chlorophyll-a and dominant planktonic components in the coastal transition zone off
Concepción, central Chile, during upwelling and non-upwelling conditions. Progress in Oceanography.
Morales, C.E., Hormazábal, S.E. & Blanco, J.L. 1999. Interannual variability in the mesoscale distribution of
the depth of the upper boundary of the oxygen minimum layer off northern Chile (18–24°S): impli-
cations for the pelagic system and biogeochemical cycling. Journal of Marine Research 57, 909–932.
Morales, C.E. & Lange, C.B. 2004 Oceanographic studies in the Humboldt Current System off Chile: an
introduction. Deep-Sea Research II 51, 2345–2348.
Morales, C.E. & Ponce, F. 1997. Parques marinos y reservas marinas en la Ley General de Pesca y Acuacultura:
desafíos para su establecimiento y aplicación. Estudios Oceanológicos 16, 19–26.
Morales, E.G. 2003. Methane hydrates in the Chilean continental margin. Biotechnology Issues for Developing
Countries 6, 80–84.
Moreno, C.A. 1995. Macroalgae as a refuge from predation for recruits of the mussel Choromytilus chorus
(Molina, 1782) in southern Chile. Journal of Experimental Marine Biology and Ecology 191, 181–193.
Moreno, C.A. 2001. Community patterns generated by human harvesting on Chilean shores: a review. Aquatic
Conservation-Marine and Freshwater Ecosystems 11, 19–30.
Moreno, C.A., Arata, J., Rubilar, P., Hucke-Gaete, R. & Robertson, G. 2006. Artisanal longline fisheries in southern
Chile: lessons to be learned to avoid incidental seabird mortality. Biological Conservation 127, 27–36.
Moreno, C.A., Asencio, G., Duarte, W.E. & Marin, V. 1998. Settlement of the muricid Concholepas conc-
holepas and its relationship with El Niño and coastal upwellings in southern Chile. Marine Ecology
Moreno, C.A., Asencio, G. & Ibáñez, S. 1993a. Settlement patterns of Concholepas concholepas (Brugiere)
(Mollusca, Muricidae) in the rocky intertidal zone of Valdivia, Chile. Revista Chilena de Historia
Natural 66, 93–101.
Moreno, C.A. & Jara, A.F. 1984. Ecological studies on fish fauna associated with Macrocystis pyrifera belt
in the north Fuegian Island, Chile. Marine Ecology Progress Series 15, 99–107.
Moreno, C.A., Lunecke, K.M. & Lépez, M.I. 1986. The response of an intertidal Concholepas concholepas
(Gastropoda) population to protection from man in southern Chile and effects on benthic sessile
assemblages. Oikos 46, 359–364.
Moreno, C.A., Reyes, A. & Asencio, G. 1993b. Habitat and movements of the recruits of Concholepas
concholepas (Mollusca, Muricidae) in the rocky intertidal of southern Chile. Journal of Experimental
Moreno, C.A. & Sutherland, J.P. 1982. Physical and biological processes in a Macrocystis pyrifera community
near Valdivia, Chile. Oecología 55, 1–6.
Moreno, R.A., Hernández, C.E., Rivadeneira, M.M., Vidal, M.A. & Rozbaczylo, N. 2006a. Patterns of endemism
in south-eastern Pacific benthic polychaetes of the Chilean coast. Journal of Biogeography 33, 750–759.
328
Moreno, R.A., Neill, P.A. & Rozbaczylo, N. 2006b. Native and non-indigenous boring polychaetes in Chile:
a threat to native and commercial mollusc species. Revista Chilena de Historia Natural 79, 263–278.
Moy, C.M., Seltzer, G.O., Rodbell, D.T. & Anderson, D.M. 2002. Variability of El Niño/Southern Oscillation
activity at millennial timescales during the Holocene epoch. Nature 420, 162–165.
Moyano, H. 1991. Bryozoa marinos chilenos. VIII: Una síntesis zoogeográfica con consideraciones sistemáti-
cas y la descripción de diez especies y dos géneros nuevos. Gayana Zoología 55, 305–389.
Mullins, H.T., Thompson, J.B., McDougall, K. & Vercoutere, T.L. 1985. Oxygen minimun zone edge effects,
evidence from the central California coastal upwelling system. Geology 13, 491–494.
Muñoz, I. 1982. Las sociedades costeras en el litoral de Arica y sus vinculaciones con la costa Peruana.
Chungará (Arica) 9, 124–151.
Muñoz, M. & Santelices, B. 1989. Determination of the distribution and abundance of the limpet Scurria
scurra on the stipes of the kelp Lessonia nigrescens in central Chile. Marine Ecology Progress Series
54, 277–285.
Muñoz, P., Lange, C.B., Gutiérrez, D., Hebbeln, D., Salamanca, M.A., Dezileau, L., Reyss, J.L. & Benninger,
L.K. 2004a. Recent sedimentation and mass accumulation rates based on Pb210 along the Perú-Chile
continental margin. Deep-Sea Research II 51, 2523–2541.
Muñoz, P., Salamanca, M.A., Neira, C. & Sellanes, J. 2004b. Nitrogen sediment fluxes in an upwelling system
off central Chile (Concepción Bay and adjacent shelf) during the 1997–1998 El Niño. Revista Chilena
Muñoz, P., Sellanes, J., Neira, C., Palma, M., Levin, L., Pantoja, S., Lange, C.B. & Salamanca, S.A. 2005.
Sedimentación v/s mezcla biológica en sedimentos del norte de Chile (Iquique, 20°S) asociados a la
zona de mínimo oxígeno (ZMO). Libro de Resúmenes, Congreso de Ciencias del Mar, Sociedad de
Ciencias del Mar de Chile, Chile, May, 172.
Muñoz, V., Hernández-González, M.C., Buschmann, A.H., Graham, M.H. & Vásquez, J.A. 2004. Variability
in per capita oogonia and sporophyte production from giant kelp gametophytes (Macrocystis pyrifera,
Phaeophyceae). Revista Chilena de Historia Natural 77, 639–647.
Murray, J. 1895. Report on the Scientific Results of the Voyage of H.M.S. Challenger. London: Eyre & Spottiswoode.
Narváez, D.A., Navarrete, S.A., Largier, J. & Vargas, C.A. 2006. Onshore advection of warm water, larval
invertebrate settlement, and relaxation of upwelling off central Chile. Marine Ecology Progress Series
309, 159–173.
Narváez, D.A., Poulin, E., Leiva, G., Hernández, E., Castilla, J.C. & Navarrete, S.A. 2004. Seasonal and
spatial variation of nearshore hydrographic conditions in central Chile. Continental Shelf Research
24, 279–292.
Narvarte, M.E., Félix-Pico, E. & Ysla-Chee, L.A. 2001. Settlement of scallop larvae on artificial collectors.
In Los Moluscos Pectínidos de Iberoamérica: Ciencia y Acuicultura, A.N. Maeda-Martínez (ed.).
Mexico City: Editorial Limusa, 173–192.
Navarrete, S.A., Broitman, B., Wieters, E.A., Finke, G.R., Venegas, R.M. & Sotomayor, A. 2002. Recruitment
of intertidal invertebrates in the southeast Pacific: interannual variability and the 1997–1998 El Niño.
Navarrete, S.A. & Castilla, J.C. 1990. Barnacle walls as mediators of intertidal mussel recruitment — effects
of patch size on the utilization of space. Marine Ecology Progress Series 68, 113–119.
Navarrete, S.A. & Castilla, J.C. 1993. Predation by Norway rats in the intertidal zone of central Chile. Marine
Navarrete, S.A. & Castilla, J.C. 2003. Experimental determination of predation intensity in an intertidal
predator guild: dominant versus subordinate prey. Oikos 100, 251–262.
Navarrete, S.A., Wieters, E.A., Broitman, B.R. & Castilla, J.C. 2005. Scales of benthic-pelagic and the intensity
of species interactions: from recruitment limitation to top-down control. Proceedings of the National
Academy of Sciences USA 102, 18,046–18,051.
Neill, P.E., Alcalde, O., Faugeron, S., Navarrete, S.A. & Correa, J.A. 2006. Invasion of Codium fragile in
northern Chile: a new threat for Gracilaria farming. Aquaculture 259, 202–210.
Neira, C., Sellanes, J., Soto, A., Gutiérrez, D. & Gallardo, V.A. 2001. Meiofauna and sedimentary organic
matter off Central Chile: response to changes caused by the 1997–1998 El Niño. Oceanologica Acta
24, 313–328.
329
MARTIN THIEL ET AL.
Neira, S. & Arancibia, H. 2004. Trophic interactions and community structure in the upwelling system off
Central Chile (33–39°S). Journal of Experimental Marine Biology and Ecology 312, 349–366.
Neira, S., Arancibia, H. & Cubillos, L. 2004. Comparative analysis of trophic structure of commercial fishery
species off Central Chile in 1992 and 1998. Ecological Modelling 172, 233–248.
Nielsen, K.J. & Navarrete, S.A. 2004. Mesoscale regulation comes from the bottom-up: intertidal interactions
between consumers and upwelling. Ecology Letters 7, 31–41.
NRC. 2001. Marine Protected Areas: Tools for Sustaining Ocean Ecosystems. Washington, D.C.: National
Academy Press.
Núñez, L. & Vásquez, J.A. 1987. Spatial distribution and trophic observations of fishes associated to a subtidal
grove of Lessonia trabeculata. Estudios Oceanológicos (Chile) 6, 79–85.
Ojeda, F.P., Labra, F.A. & Muñoz, A.A. 2000. Biogeographic patterns of Chilean littoral fishes. Revista Chilena
Ojeda, F.P. & Santelices, B. 1984. Ecological dominance of Lessonia nigrescens (Phaeophyta) in Central
Chile. Marine Ecology Progress Series 19, 83–91.
Oliger, P. & Santelices, B. 1981. Physiological ecology studies on Chilean Gelidiales. Journal of Experimental
Olivares, G. 2001. Mecanismos de interacción físico-químicas en una zona de surgencia costera: retención
de larvas y cierre del ciclo de vida de Euphausia mucronata. M.Sc. thesis, Universidad de Chile,
Santiago, Chile.
Olivieri, R.A. & Chavez, F.P. 2000. A model of plankton dynamics for the coastal upwelling system of
Monterey Bay, California. Deep-Sea Research II 47, 1077–1106.
Ortiz, M. 2003. Qualitative modelling of the kelp forest of Lessonia nigrescens Bory (Laminariales: Phaeo-
phyta) in eulittoral marine ecosystems of the south-east Pacific: an approach to management plan
assessment. Aquaculture 220, 423–436.
Ortiz, M., Jesse, S., Stotz, W. & Wolff, M. 2003. Feeding behaviour of the asteroid Meyenaster gelatinosus
in response to changes in abundance of the scallop Argopecten purpuratus in northern Chile. Archiv
für Hydrobiologie 157, 213–225.
Ortiz, M. & Stotz, W. 1996. Distribución de juveniles de asentamiento reciente de Mesodesma donacium
(Lamarck, 1818) (Mollusca: Bivalvia: Mesodesmatidae) en tres bahías de la IV Región: variables
físicas, químicas y biológicas que le caracterizan. Biología Pesquera 25, 27–40.
Ortlieb, L. 1995. Paleoclimas cuaternarios en el norte grande de Chile. In Cambios Cuaternarios en América
del Sur, J. Argollo & P. Mourguiart (eds). La Paz, Bolivia: ORSTOM-Bolivia, 225–246.
Ortlieb, L., Escribano, R., Follegati, R., Zuniga, O., Kong, I., Rodriguez, L., Valdes, J., Guzmán, N. &
Iratchet, P. 2000. Recording of ocean-climate changes during the last 2,000 years in a hypoxic marine
environment off northern Chile (23°S). Revista Chilena de Historia Natural 73, 221–242.
Ortlieb, L., Guzmán, N. & Candia, M. 1994. Moluscos litorales del Pleistoceno superior en el área de
Antofagasta, Chile: primeras determinaciones e indicaciones paleoceanográficas. Estudios Ocean-
ológicos 13, 57–63.
Oyarzún, C. 1998. El Jurel, ¿de qué estamos hablando? In Biología y Ecología del Jurel en Aguas Chilenas,
D. Arcos (ed.). Talcahuano, Chile: Instituto de Investigación Pesquera, 37–45.
Pacheco, A. & Stotz, W.B. 2006. Will providing a filamentous substratum in the water column and shell litter
on the bottom increase settlement and post-larval survival of the scallop Argopecten purpuratus?
Pagès, F., González, H.E., Ramon, M., Sobarzo, M. & Gili, J.M. 2001. Gelatinous zooplankton assemblages
associated with water masses in the Humboldt Current System, and potential predatory impact by
Bassia bassensis (Siphonophora: Calycophorae). Marine Ecology Progress Series 210, 13–24.
Paine, R.T., Castilla, J.C. & Cancino, J.M. 1985. Perturbation and recovery patterns of starfish-dominated
intertidal assemblages in Chile, New Zealand, and Washington State. American Naturalist 125,
679–691.
Palma, A.T. & Ojeda, F.P. 2002. Abundance, distribution and feeding patterns of a temperate reef fish in
subtidal environments of the Chilean coast: the importance of understory algal turf. Revista Chilena
330
Palma, A.T., Pardo, L.M., Veas, R., Cartes, C., Silva, M., Manríquez, K., Díaz, A., Muñoz, C. & Ojeda, F.P.
2006. Coastal brachyuran decapods: settlement and recruitment under contrasting coastal geometry
conditions. Marine Ecology Progress Series 316, 139–153.
Palma, M., Quiroga, E., Gallardo, V.A., Arntz, W., Gerdes, D., Schneider, W. & Hebbeln, D. 2005. Macro-
benthic animal assemblages of the continental margin off Chile (22° to 42°S). Journal of the Marine
Palma, S. 1994. Distribución y abundancia de larvas de langostino colorado Pleuroncodes monodon frente a
la costa de Concepción, Chile. Investigaciones Marinas, Valparaíso 22, 13–29.
Palma, S. & Arana, P. 1997. Aspectos reproductivos del langostino colorado (Pleuroncodes monodon H. Milne
Edwards, 1837) frente a la costa de Concepción, Chile. Investigaciones Marinas, Valparaíso 25,
203–221.
Palma, W., Escribano, R. & Rosales, S.A. 2006. Modeling study of seasonal and inter-annual variability of
circulation in the coastal upwelling site of the El Loa River off northern Chile. Estuarine Coastal and
Palmer, A.R. & Strathmann, R.R. 1981. Scale of dispersal in varying environments and its implications for
life histories of marine invertebrates. Oecologia 48, 308–318.
Palumbi, S.R. 1995. Using genetics as an indirect estimator of larval dispersal. In Ecology of Marine Inver-
tebrate Larvae, L.R. McEdward (ed.). Boca Raton, FL: CRC Press, 369–387.
Palumbi, S.R. 2003. Population genetics, demographic connectivity, and the design of marine reserves.
Ecological Applications 13, S146–S158.
Pantoja, S., Sepulveda, J. & González, H.E. 2004. Decomposition of sinking proteinaceous material during
fall in the oxygen minimum zone off northern Chile. Deep-Sea Research I 51, 55–70.
Pauchard, A. & Villarroel, P. 2002. Protected areas in Chile: history, current status, and challenges. Natural
Areas Journal 22, 318–330.
Pauly, D., Muck, P., Mendo, J. & Tsukayama, I. (eds) 1989. The Peruvian Upwelling Ecosystem: Dynamics
and Interactions. Callao, Peru: ICLARM Conference Proceedings.
Pavez, M.A., Castro, L.R. & González, H.E. 2006. Across-shelf predatory effect of Pleurobrachia bachei
(Ctenophora) on the small-copepod community in the coastal upwelling zone off northern Chile
(23°S). Journal of Plankton Research 28, 115–129.
Pechenik, J.A. 1986. Field evidence for delayed metamorphosis of larval gastropod: Crepidula fornicata,
C. plana, and Bittium alternatum. Journal of Experimental Marine Biology and Ecology 97, 313–319.
Pechenik, J.A. 1999. On the benefits of being a larva … or not. In Aquatic Life Cycle Strategies, M. Whitfield
et al. (eds). Plymouth, U.K.: Marine Biological Association of the United Kingdom, 97–104.
Peña, T.S., Johst, K., Grimm, V., Arntz, W.E. & Tarazona, J. 2006. Disentangling the effects of El Niño on a
population of the polychaete Sigambra bassi in the Bay of Ancón, Peru. Advances in Geosciences 6,
161–166.
Peñalver, E. 2004. Dinámica de la capa superior de la zona costera frente a Coquimbo y su relación con la
distribución de nutrientes y clorofila. M.Sc. thesis, Universidad Católica del Norte, Coquimbo, Chile.
Penven, P. 2003. ROMSTOOLS User’s Guide, Tech Report. Inst. de Rech. pour le Dev., Paris, France. Available
HTTP: http://www.brest.ird.fr/Roms_tools. Accessed 21 February 2007.
Pérez, E.P. 2003. Análisis bioeconómico de la pesquería de crustáceos en la plataforma centro-norte de Chile.
Ph.D. Thesis, Centro de Investigación y Estudios Avanzados, Unidad de Mérida, Mérida, Mexico.
Pérez, E.P. 2005. Un modelo simple para describir la dinámica de la biomasa del camarón nailon Heterocarpus
reedi en Coquimbo, Chile. Investigaciones Marinas, Valparaíso 33, 131–142.
Pérez, E.P. & Stotz, W. 1992. Comparaciones múltiples de parámetros gravimétricos entre poblaciones
submareales de Concholepas concholepas (Bruguiére, 1789) en el norte de Chile. Revista de Biología
Marina, Valparaíso (Chile) 27, 175–186.
Peters, A.F. & Breeman, A.M. 1993. Temperature tolerance and latitudinal range of brown algae from temperate
Pacific South America. Marine Biology 115, 143–150.
Peterson, W.T., Arcos, D.F., McManus, G.B., Dam, H., Bellantoni, D., Johnson, T. & Tiselius, P. 1988. The
nearshore zone during coastal upwelling daily variability and coupling between primary and secondary
production off central Chile. Progress in Oceanography 20, 1–40.
331
MARTIN THIEL ET AL.
Phillips, N.E. 2002. Benthic-pelagic coupling in intertidal communities: integrating across the life cycle of
filter-feeders marine invertebrates. Ph.D. thesis, University of California, Santa Barbara.
Phillips, R.A., Silk, J.R, Croxall, J.P. & Afanasyev, V. 2006. Year-round distribution of white-chinned petrels from
South Georgia: relationships with oceanography and fisheries. Biological Conservation 129, 336–347.
Pineda, J. 1991. Predictable upwelling and the shoreward transport of planktonic larvae by internal tidal bores.
Science 253, 548–551.
Pineda, J. 1994a. Internal tidal bores in the nearshore warm water fronts, seaward gravity currents and the
onshore transport of neustonic larvae. Journal of Marine Research 52, 427–458.
Pineda, J. 1994b. Spatial and temporal patterns in barnacle settlement rate along a southern California rocky
shore. Marine Ecology Progress Series 107, 125–138.
Piñones, A., Valle-Levinson, A., Narváez, D.A., Vargas, C.A., Navarrete, S.A., Yuras, G. & Castilla, J.C. 2005.
Wind-induced diurnal variability in river plume motion. Estuarine Coastal and Shelf Science 65, 513–525.
Pizarro, A. & Santelices, B. 1993. Environmental variation and large-scale Gracilaria production. Hydrobio-
logia 260/261, 357–363.
Pizarro, G., Iriarte, J.L. & Montecino, V. 2002. Mesoscale primary production and bio-optical variability off
Antofagasta (23–24°S) during the transition to El Niño 1997–1998. Revista Chilena de Historia
Natural 75, 201–215.
Pizarro, J., Palma, W. & Flores, C. 1998. Validación de la hipótesis de mortalidad por inanición en larvas de
Engraulis ringens Jenyns, 1842 (Pisces: Engraulidae), utilizando el criterio histológico en un área de
surgencia costera en el norte de Chile. Investigaciones Marinas, Valparaíso 26, 97–108.
Polis, G.A. & Hurd, S.D. 1995. Extraordinarily high spider densities on islands — flow of energy from the
marine to terrestrial food webs and the absence of predation. Proceedings of the National Academy
of Sciences USA 92, 4382–4386.
Polis, G.A. & Hurd, S.D. 1996. Linking marine and terrestrial food webs: allochthonous input from the ocean
supports high secondary productivity on small islands and coastal land communities. American
Polis, G.A., Hurd, S.D., Jackson, C.T. & Pinero, F.S. 1997. El Niño effects on the dynamics and control of
an island ecosystem in the Gulf of California. Ecology 78, 1884–1897.
Pomeroy, R.S., Parks, J.E. & Watson, L.M. 2004. How Is Your MPA Doing? A Guidebook of Natural and Social
Indicators for Evaluating Marine Protected Area Management Effectiveness. Gland, Switzerland: IUCN.
Poulin, E., Cárdenas, L., Hernández, C.E., Kornfield, I. & Ojeda, F.P. 2004. Resolution of the taxonomic status
of Chilean and Californian jack mackerels using mitochondrial DNA sequence. Journal of Fish Biology
65, 1160–1164.
Poulin, E., Palma, A.T., Leiva, G., Hernández, E., Martínez, P., Navarrete, S.A. & Castilla, J.C. 2002b. Temporal
and spatial variation in the distribution of epineustonic competent larvae of Concholepas concholepas
along the central coast of Chile. Marine Ecology Progress Series 229, 95–104.
Poulin, E., Palma, A.T., Leiva, G., Narváez, D., Pacheco, R., Navarrete, S.A. & Castilla, J.C. 2002a. Avoiding
offshore transport of competent larvae during upwelling events: the case of the gastropod Concholepas
concholepas in Central Chile. Limnology and Oceanography 47, 1248–1255.
Prado, L. & Castilla, A.C. 2006. The bioengineer Perumytilus purpuratus (Mollusca: Bivalvia) in central Chile:
biodiversity, habitat structural complexity and environmental heterogeneity. Journal of the Marine
Pulgar, J.M., Ojeda, F.P. & Bozinovic, F. 2006. Intraspecific geographic and seasonal physiological variability
in an intertidal fish, Girella laevifrons, along a climatic gradient. Journal of Fish Biology 68, 975–981.
Quijón, P., Jaramillo, E. & Contreras, H. 2001. Distribution and habitat structure of Ocypode gaudichaudii
H. Milne Edwards & Lucas, 1843, in sandy beaches of northern Chile. Crustaceana 74, 91–103.
Quiroga, E., Soto, R. & Rozbaczylo, N. 1999. Los poliquetos espiónidos (Polychaeta: Spionidae) y su
importancia en la estructura de una comunidad: un caso de estudio en bahía Iquique, norte de Chile
(20°11′S, 70°10′W). Gayana 63, 1–16.
Quiroz, J.C., Wiff, R. & Montenegro, C. 2005. Factores que afectan las tasas de captura de langostino amarillo
(Cervimunida johni) en la zona norte de Chile. Investigaciones Marinas, Valparaíso 33, 43–55.
Radashevsky, V.I. & Olivares, C. 2005. Polydora uncinata (Polychaeta: Spionidae) in Chile: an accidental
transportation across the Pacific. Biological Invasions 7, 489–496.
332
Ramírez, A. 2005a. La transformación epistemológica contemporánea. De la Unidad a la dispersión. Santiago,

Chile: Editorial Universitaria.
Ramírez, A. 2005b. Estudio del capital natural y los servicios ecosistémicos asociados a las actividades de
observación de la naturaleza, en la reserva Pingüino de Humboldt, Región de Coquimbo, Chile.
Honors thesis, Universidad de Chile, Santiago, Chile.
Ramorino, L. & Muñiz, L. 1970. Estudio cuantitativo general sobre la fauna de fondo de la bahía de Mejillones.
Revista de Biología Marina, Valparaíso (Chile) 14, 79–93.
Ramorino, L.M. 1968. Pelecypoda del fondo de la Bahía de Valparaíso. Revista de Biología Marina, Valparaíso
(Chile) 13, 176–285.
Ramos, M., Pizarro, O., Bravo, L. & Dewitte, B. 2006. Seasonal variability of the permanent thermocline off
northern Chile. Geophysical Research Letters 33, L09608, DOI:10.1029/2006GL025882.
Reed, D.C., Amsler, C.D. & Ebeling, A.W. 1992. Dispersal in kelps — factors affecting spore swimming and
competence. Ecology 73, 1577–1585.
Rendell, L., Whitehead, H. & Escribano, R. 2004. Sperm whale habitat use and foraging success off northern
Chile: evidence of ecological links between coastal and pelagic systems. Marine Ecology Progress
Series 275, 289–295.
Rivadeneira, M.M. 2005. Macroecología evolutiva de los bivalvos marinos de la costa pacífica de Sudamérica.
Ph.D. thesis, Pontificia Universidad Católica de Chile, Santiago, Chile.
Rivadeneira, M.M. & Fernández, M. 2005. Shifts in southern endpoints of distribution in rocky intertidal
species along the south-eastern Pacific coast. Journal of Biogeography 32, 203–209.
Rivera, G. 1992. Taxonomía y fenología reproductiva de Gastroclonium parvum (Hollenberg) Chang y Xia
(Rhodymeniales, Champiaceae). Honors thesis, Universidad Católica del Norte, Coquimbo, Chile.
Roa, R., Gallardo, V.A., Ernst, B., Baltazar, M., Cañete, J.I. & Enríquez-Briones, S. 1995. Nursery ground,
age structure and abundance of juvenile squat lobster Pleuroncodes monodon on the continental shelf
off central Chile. Marine Ecology Progress Series 116, 47–54.
Roa, R. & Tapia, F. 1998. Spatial differences in growth and sexual maturity between branches of a large
population of the squat lobster Pleuroncodes monodon. Marine Ecology Progress Series 167, 185–196.
Roberts, C.M., Branch, G., Bustamante, R.H., Castilla, J.C., Dugan, J., Halpern, B.S., Lafferty, K.D., Leslie, H.,
Lubchenco, J., McArdle, D., Ruckelshaus, M. & Warner, R.R. 2003. Application of ecological criteria
in selecting marine reserves and developing reserve networks. Ecological Applications 13, S215–S228.
Rodríguez, L., Escribano, R., Grone, G., Irribarren, C. & Castro, H. 1996. Ecología del fitoplancton en la
Bahía de Antofagasta (23°S), Chile. Revista de Biología Marina, Valparaíso (Chile) 31, 65–80.
Rodríguez, S.R. 2003. Consumption of drift kelp by intertidal populations of the sea urchin Tetrapygus niger
on the central Chilean coast: possible consequences at different ecological levels. Marine Ecology
Rodríguez-Graña, L. & Castro, L.R. 2003. Ichthyoplankton distribution off the Peninsula de Mejillones, Chile
(23°S, 71°W), under variable hydrographic conditions during the austral summer and winter of the
1997 El Niño. Hydrobiologia 501, 59–73.
Rodríguez-Graña, L., Castro, L., Loureiro, M., González, H.E. & Calliari, D. 2005. Feeding ecology of
dominant larval myctophids in an upwelling area of the Humboldt Current. Marine Ecology Progress
Series 290, 119–134.
Rojas, P.M., Escribano, R. & Marín, V.H. 2002. Fish larvae distribution off Mejillones Peninsula (northern
Chile) during a coastal upwelling event in Spring 1999: interactions with the cold upwelling plume.
Fisheries Oceanography 11, 233–244.
Ronan, C.E. 2002. Juan Ignacio Molina: The World’s Window on Chile. New York: Peter Lang Publishing.
Rosenberg, R., Arntz, W., Chumán de Flores, E., Flores, L.A., Carbajal, G., Finger, I. & Tarazona, J. 1983.
Benthos biomass and oxygen deficiency in the Peruvian upwelling system. Journal of Marine Research
41, 263–279.
Roughgarden, J., Gaines, S. & Possingham, H. 1988. Recruitment dynamics in complex life cycles. Science
241, 1460–1466.
Roughgarden, J., Pennington, J.T., Stoner, D., Alexander, S. & Miller, K. 1991. Collisions of upwelling fronts
with the intertidal zone — the cause of recruitment pulses in barnacle populations of central California.
Acta Oecologica 12, 35–51.
333
MARTIN THIEL ET AL.
Rovira, J. 2006. Políticas Públicas para la Conservación y Uso Sostenible de La Biodiversidad: Un Aporte
para la Convivencia entre Minería y Conservación. Santiago, Chile: Publicaciones de Sonami Chile.
Quebecor World Chile S.A.
Roy, C. 1998. An upwelling-induced retention area off Senegal: a mechanism to link upwelling and retention
processes. South African Journal of Marine Sciences 19, 89–98.
Ruiz-Tagle, N., Fernández, M. & Pörtner, H.O. 2002. Full time mothers: daily rhythms in brooding and nonbrood-
ing behaviors of brachyuran crabs. Journal of Experimental Marine Biology and Ecology 276, 31–47.
Rutllant, J.A., Fuenzalida, H., Torres, R. & Figueroa, D. 1998. Land-air-sea interaction at the Antofagasta
region (Chile, 23°S). The DICLIMA experiment. Revista Chilena de Historia Natural 71, 405–427.
Rutllant, J.A., Masotti, I., Calderon, J. & Vega, S.A. 2004b. A comparison of spring coastal upwelling off
central Chile at the extremes of the 1996–1997 ENSO cycle. Continental Shelf Research 24, 773–787.
Rutllant, J.A. & Montecino, V. 2002. Multiscale upwelling forcing cycles and biological response off north-
Rutllant, J.A., Rosenbluth, B. & Hormazábal, S. 2004a. Intraseasonal variability of wind-forced coastal
upwelling off central Chile (30°S). Continental Shelf Research 24, 789–804.
Ryther, J.H. 1969. Photosynthesis and fish production in the sea. Science 166, 72–76.
Sabat, P., Fariña, J.M. & Soto-Gamboa, M. 2003. Terrestrial birds living on marine environments: does dietary
composition of Cinclodes nigrofumosus (Passeriformes: Furnariidae) predict their osmotic load?
Revista Chilena de Historia Natural 76, 335–343.
Sahli, J.M. 2006. Distribución espacial del género Nassarius (Gastropoda: Prosobranchia) en el bentos de la
Bahía de La Herradura, Coquimbo. Honors thesis, Universidad Católica del Norte, Coquimbo, Chile.
Saito, M. 2005. Cobalt, iron, and manganese biogeochemistry in the equatorial Pacific and recovery experi-
ments using the MgOH2 precipitation ICP-MS method. Abstracts of Papers of the American Chemical
Society 229, 894.
Sala, E., Aburto-Oropeza, O., Paredes, G., Parra, I., Barrera, J.C. & Dayton, P.K. 2002. A general model for
designing networks of marine reserves. Science 298, 1991–1993.
Salen-Picard, C., Darnaude, A.M., Arlhac, D. & Harmelin-Vivien, M.L. 2002. Fluctuations of macrobenthic
populations: a link between climate driven river run off and sole fishery yields in the Gulf of Lions.
Oecologia 133, 380–388.
Sánchez, M., Castilla, J.C. & Mena, O. 1982. Variaciones verano-invierno de la macrofauna de arena en playa
Morrillos (norte chico, Chile). Studies on Neotropical Fauna and Environment 17, 31–49.
Sanchez-Pinero, F. & Polis, G.A. 2000. Bottom-up dynamics of allochthonous input: direct and indirect effects
of seabirds on islands. Ecology 81, 3117–3132.
Sanders, H. 1969. Benthic marine diversity and stability-time hypothesis. Brookhaven Symposium of Biology
22, 71–81.
Sandweiss, D.H. 2003. Terminal pleistocene through mid-holocene archaeological sites as paleoclimatic
archives for the Peruvian coast. Palaeogeography Palaeoclimatology Palaeoecology 194, 23–40.
Sandweiss, D.H., Maasch, K.A., Burger, R.L., Richardson, J.B., Rollins, H.B. & Clement, A. 2001. Variation
in Holocene El Niño frequencies: climate records and cultural consequences in ancient Perú. Geology
29, 603–606.
Sandweiss, D.H., McInnis, H., Burger, R.L., Cano, A., Ojeda, B., Paredes, R., Sandweiss, M.D. & Glascock,
M.D. 1998. Quebrada Jaguay: early South American maritime adaptations. Science 281, 1830–1832.
Santelices, B. 1980. Phytogeographic characterization of the temperate coast of Pacific South America.
Santelices, B. 1989. Algas Marinas de Chile. Santiago, Chile: Ediciones Universidad Católica de Chile.
Santelices, B. 1990a. Patterns of reproduction, dispersal and recruitment in seaweeds. Oceanography and
Marine Biology: An Annual Review 28, 177–276.
Santelices, B. 1990b. Patterns of organizations of intertidal and shallow subtidal vegetation in wave exposed
habitats of central Chile. Hydrobiologia 192, 35–57.
Santelices, B. 1991. Littoral and sublittoral communities of continental Chile. In Ecosystems of the World.
Intertidal and Littoral Ecosystems, D. Goodhall (ed.). Amsterdam: Elsevier, 347–369.
Santelices, B., Aedo, D. & Hoffmann, A. 2002. Banks of microscopic forms and survival to darkness of
propagules and microscopic stages of macroalgae. Revista Chilena de Historia Natural 75, 547–555.
334
Santelices, B., Castilla, J.C., Cancino, J. & Schmiede, P. 1980. Comparative ecology of Lessonia nigrescens
and Durvillaea antarctica (Phaeophyta) in central Chile. Marine Biology 59, 119–132.
Santelices, B. & Correa, J.A. 1985. Differential survival of macroalgae to digestion by intertidal herbivore
molluscs. Journal of Experimental Marine Biology and Ecology 88, 183–191.
Santelices, B., Correa, J.A., Aedo, D., Flores, V., Hormázabal, M. & Sánchez, P. 1999. Convergent biological
processes in coalescing Rhodophyta. Journal of Phycology 35, 1127–1149.
Santelices, B., Correa, J.A., Hormazábal, M. & Flores, V. 2003. Contact responses between spores and
sporelings of different species, karyological phases and cystocarps of coalescing Rhodophyta. Marine
Biology 143, 381–392.
Santelices, B. & Marquet, P.A. 1998. Seaweeds, latitudinal diversity patterns, and Rapoport’s rule. Diversity
and Distribution 4, 71–75.
Santelices, B. & Meneses, I. 2000. A reassessment of the phytogeographic characterization of temperate Pacific
South America. Revista Chilena de Historia Natural 73, 605–614.
Santelices, B. & Norambuena, R. 1987. A harvesting strategy for Iridaea laminarioides in Central Chile.
Hydrobiologia 151/152, 329–333.
Santelices, B. & Ojeda, F.P. 1984. Recruitment, growth and survival of Lessonia nigrescens (Phaeophyta) at
various tidal levels in exposed habitats of central Chile. Marine Ecology Progress Series 19, 73–82.
Santelices, B. & Payá, I. 1989. Digestion survival of algae. Qualitative and quantitative comparisons between
spores and fecal pellets. Journal of Phycology 25, 693–699.
Santelices, B., Vásquez, J.A., Ohme, U. & Fonck, E. 1984. Managing wild crops of Gracilaria in central
Chile. Hydrobiologia 116/117, 77–89.
Santibáñez, J. 1944. La Corriente de Humboldt en las Costas de Chile. Valparaíso, Chile: Liga Marítima de Chile.
Santoro, C.M., Arriaza, B.T., Standen, V.G. & Marquet, P.A. 2005. People of the coastal Atacama desert:
living between sand dunes and waves of the Pacific Ocean. In Desert Peoples: Archeological Per-
spectives, P. Veth et al. (eds). Oxford, U.K.: Blackwell Publishing, 243–260.
Sayre, R., Roca, E., Sedaghatkish, G., Young, B., Keel, S. & Sheppard, S. 2000. Nature in Focus — Rapid
Ecological Assessment. Washington, D.C.: Island Press.
Secretariat of the Convention on Biological Diversity. 2004. Technical advice on the establishment and
management of a national system of marine and coastal protected areas, SCBD. CBD Technical Series
13, 1–40.
Scheltema, R.S. 1986. On dispersal and planktonic larvae of benthic invertebrates: an eclectic overview and
summary of problems. Bulletin of Marine Science 39, 290–322.
Schlatter, R.P. 1984. Status and conservation of seabirds in Chile. In Status and Conservation of the World’s
Seabirds, J.P. Croxall & R.W. Schreiber (eds). Cambridge, U.K.: ICBP Technical Publication, 261–269.
Schlatter, R.P. 1987. Conocimiento y situación de la ornitofauna en las islas oceánicas chilenas. In Islas
Oceánicas Chilenas: Conocimiento Científico y Necesidades de Investigaciones, J.C. Castilla (ed.).
Santiago, Chile: Ediciones Universidad Católica de Chile, 271–285.
Schlatter, R.P. & Simeone, A. 1999. Estado del conocimiento y conservación de las aves en mares chilenos.
Estudios Oceanológicos 18, 25–33.
Schmalfuss, H. & Vergara, K. 2000. The isopod genus Tylos (Oniscidae: Tylidae) in Chile, with bibliographies
of all described species of the genus. Stuttgarter Beiträge zur Naturkunde, Serie A 612, 1–42.
Schmidt-Nielsen, K. 1997. Animal Physiology. Cambridge: Cambridge University Press.
Schulz, H.N., Strotmann, B., Gallardo, V.A. & Jorgensen, B.B. 2000. Population study of the filamentous
sulfur bacteria Thioploca spp. off the Bay of Concepción, Chile. Marine Ecology Progress Series
200, 117–126.
Sebens, K.P. & Paine, R.T. 1979. Biogeography of anthozoans along the west coast of South America: habitat,
disturbance, and prey availability. Proceedings of the International Symposium of Marine Biogeog-
raphy and Evolution in the Southern Hemisphere 2, 219–237.
Sellanes, J. & Krylova, E. 2005. A new species of Calyptogena (Bivalvia: Vesicomyidae) from a recently
discovered methane seepage area off Concepción Bay, Chile (~36°S). Journal of the Marine Biological
Sellanes, J. & Neira, C. 2006. ENSO as a natural experiment to understand environmental control of meiofaunal
community structure. Marine Ecology 27, 31–43.
335
MARTIN THIEL ET AL.
Sellanes, J., Neira, C. & Quiroga, E. 2003. Composition, structure and energy flux of the meiobenthos off
Sellanes, J., Quiroga, E., Neira, C. & Gutierrez, D. 2007. Changes of macrobenthos composition under different
ENSO cycle conditions on the continental shelf off central Chile. Continental Shelf Research, 27,
1002–1016.
Sellanes, J., Quiroga, E. & Gallardo, V.A. 2004. First direct evidence of methane seepage and associated
chemosynthetic communities in the bathyal zone off Chile. Journal of the Marine Biological Associ-
ation of the United Kingdom 84, 1065–1066.
Semenov, V.N. 1977. Biogeographical latitudinal-zonal nomenclatura of coastal marine biota. Oceanology 17,
90–96.
Sepúlveda, R., Cancino, J.M. & Thiel, M. 2003a. The peracarid epifauna associated with the ascidian Pyura
chilensis (Molina, 1782) (Ascidiacea: Pyuridae). Journal of Natural History 37, 1555–1569.
Sepúlveda, R., Moreno, R.A. & Carrasco, F.D. 2003b. Diversidad de macroinvertebrados asociados a arrecífes
de Phragmatopoma moerchi Kinberg, 1867 (Polychaeta: Sabellariidae) en el intermareal rocoso de
Cocholgüe, Chile. Gayana 67, 45–54.
SERNAPESCA. 2005. Anuario Estadístico de Pesca del Servicio Nacional de Pesca. Valparaíso, Chile:
Publicación del Departamento de Sistemas de Información y Estadísticas Pesqueras, SERNAPESCA.
Available HTTP: http://www.sernapesca.cl./index.php. Accessed 21 February 2007.
Shaffer, G., Hormázabal, S., Pizarro, O. & Salinas, S. 1999. Seasonal and interannual variability of currents
and temperature off central Chile. Journal of Geophysical Research-Oceans 104, 29,951–29,961.
Shaffer, G., Pizarro, O., Djurfeldt, L., Salinas, S. & Rutllant, J. 1997. Circulation and low frequency variability
near the Chilean coast: remotely forced fluctuations during the 1991–1992 El Niño. Journal of Physical
Shaffer, G., Salinas, S., Pizarro, O., Vega, A. & Hormazábal, S. 1995. Currents in the deep-ocean off Chile
(30°S). Deep-Sea Research I 42, 425–436.
Shanks, A.L. 1983. Surface slicks associated with tidally forced internal waves may transport pelagic larvae
of benthic invertebrates and fishes shoreward. Marine Ecology Progress Series 13, 311–315.
Shanks, A.L. 1995. Mechanisms of cross-shelf dispersal of larval invertebrates. In Ecology of Marine Inver-
tebrate Larvae, L. McEdward (ed.). Boca Raton, Florida: CRC Press, 232–267.
Shanks, A.L. 2002. Internal tides and the biology of continental shelf waters. In The Oceanography and
Ecology of the Nearshore and Bays in Chile, J.C. Castilla & J.L. Largier (eds). Santiago, Chile:
Shanks, A.L., Grantham, B.A. & Carr, M.H. 2003a. Propagule dispersal distance and the size and spacing of
marine reserves. Ecological Applications 13, S159–S169.
Shanks, A.L., McCulloch, A. & Miller, J. 2003b. Topographically generated fronts, very nearshore oceanography
and the distribution of larval invertebrates and holoplankters Journal of Plankton Research 25, 1251–1277.
Shchepetkin, A.F. & McWilliams, J.C. 2005. The regional oceanic modeling system (ROMS): a split-explicit,
free-surface, topography-following-coordinate oceanic model. Ocean Modelling 9, 347–404.
Shears, N.T. & Babcock, R.C. 2003. Continuing trophic cascade effects after 25 years of no-take marine
reserve protection. Marine Ecology Progress Series 246, 1–16.
Siegel, D.A., Kinlan, B.P., Gaylord, B. & Gaines, S.D. 2003. Lagrangian descriptions of marine larval
dispersion. Marine Ecology Progress Series 260, 83–96.
Sielfeld, W. 1999. Estado del conocimiento sobre conservación y preservación de Otaria flavescens (Shaw,
1800) y Arctocephalus australis (Zimmermann, 1783) en las costas de Chile. Estudios Oceanologicos
18, 81–96.
Sielfeld, W., Amado, N., Peredo, R., Vargas, M., Guerra, C., Malinarich, A., Acuña, E., Cerda, G., Bolvaran,
A., Durán, R., Aguayo, A., Sepúlveda, M., Palma, F., Veloso, X., Guerra, Y. & Galáz, J. 1997.
Monitoreo de la pesquería y censo del lobo marino común en el litoral de la I-IV Regiones. Proyecto
Fondo de Investigación Pesquera FIP. No. 95-28. Universidad Arturo Prat, Iquique, Chile.
Sielfeld, W. & Castilla, J.C. 1999. Estado de conservación y conocimiento de las nutrias en Chile. Estudios
Oceanológicos 18, 69–79.
Sielfeld, W. & Vargas, M. 1996. Composición y estructura de la ictiofauna demersal en la zona norte de Chile.
Investigaciones Marinas, Valparaíso 24, 3–17.
336
Sielfeld, W., Vargas, M., Berríos, V. & Aguirre, G. 2002. Warm ENSO events and their effects on the coastal
fish fauna of northern Chile. Investigaciones Marinas, Valparaíso 30, 122–124.
Silva, C., Yáñez, E., Barbieri, M.A. & Nieto, K. 2003. Asociaciones entre la pesquería de pequeños pelágicos,
la clorofila a y la temperatura superficial del mar en la zona norte de Chile. In Actividad Pesquera y
de Acuicultura en Chile, E. Yáñez (ed.). Valparaíso, Chile: Escuela de Ciencias del Mar, Universidad
Católica de Valparaíso, 157–178.
Silva, N. & Neshyba, S. 1979. On the southern-most extension of the Perú-Chile undercurrent. Deep-Sea
Research I 26, 1387–1393.
Simeone, A., Araya, B., Bernal, M., Diebold, E.N., Grzybowski, K., Michaels, M., Teare, J.A., Wallace, R.S.
& Willis, M.J. 2002. Oceanographic and climatic factors influencing breeding and colony attendance
patterns of Humboldt penguins Spheniscus humboldti in central Chile. Marine Ecology Progress Series
227, 43–50.
Simeone, A., Bernal, M. & Meza, J. 1999. Incidental mortality of Humboldt penguins Spheniscus humboldti
in gill nets, central Chile. Marine Ornithology 27, 157–161.
Simeone, A., Luna-Jorquera, G., Bernal, M., Garthe, S., Sepúlveda, F., Villablanca, R., Ellenberg, U., Contreras,
M., Muñoz, J. & Ponce, T. 2003. Breeding distribution and abundance of seabirds on islands off north-
Smith, C.R., Mincks, S. & DeMaster, D.J. 2006. A synthesis of bentho-pelagic coupling on the Antarctic
shelf: food banks, ecosystem inertia and global climate change. Deep-Sea Research II 53, 875–894.
Sobarzo, M. & Figueroa, D. 2001. The physical structure of a cold filament in a Chilean upwelling zone
(Peninsula de Mejillones, Chile, 23°S). Deep-Sea Research I 48, 2699–2726.
Sobel, J.A. & Dahlgren, C.P. 2004. Marine Reserves: A Guide to Science, Design And Use. Washington, D.C.:
Island Press.
Sobolev, I. 2000. Effect of column ozone on the variability of biologically effective UV radiation at high
southern latitudes. Photochemistry and Photobiology 72, 753–765.
Somero, G.N. 2002. Thermal physiology and vertical zonation of intertidal animals: optima, limits, and costs
of living. Integrative and Comparative Biology 42, 780–789.
Sotka, E.E. & Palumbi, S.R. 2006. The use of genetic clines to estimate dispersal distances of marine larvae.
Ecology 87, 1094–1103.
Soto, R. 1985. Efectos del fenómeno de El Niño 1982–1983 en ecosistemas de la I Región. Investigaciones
Pesqueras (Chile) 32, 199–206.
Spicer, J.L. & Gaston, K.J. 1999. Physiological Diversity and Its Ecological Implications. Oxford, U.K.:
Blackwell Science.
Stepien, C.A. & Rosenblatt, R.H. 1996. Genetic divergence in antitropical pelagic marine fishes (Trachurus,
Merluccius, and Scomber) between North and South America. Copeia 1996, 586–598.
Stillman, J.H. 2002. Causes and consequences of thermal tolerance limits in rocky intertidal porcelain crabs,
genus Petrolisthes. Integrative and Comparative Biology 42, 790–796.
Stillman, J.H. & Somero, G.N. 1996. Adaptation to temperature stress and aerial exposure in congeneric
species of intertidal porcelain crabs (genus Petrolisthes): correlation of physiology, biochemistry and
morphology with vertical distribution. Journal of Experimental Biology 199, 1845–1855.
Stillman, J.H. & Somero, G.N. 2000. A comparative analysis of the upper thermal tolerance limits of eastern
Pacific porcelain crabs, genus Petrolisthes: influences of latitude, vertical zonation, acclimation, and
phylogeny. Physiological and Biochemical Zoology 73, 200–208.
Stone, C.J. 1989. A comparison of algal diets for cirripede nauplii. Journal of Experimental Marine Biology
and Ecology 132, 17–40.
Stotz, W. 1997. Las áreas de manejo en la ley de pesca y acuicultura: primeras experiencias y evaluación de
la utilidad de esta herramienta para el recurso loco. Estudios Oceanológicos 16, 67–86.
Stotz, W. 2000. When aquaculture restores and replaces an overfished stock: is the conservation of the species
assured? The case of the scallop Argopecten purpuratus in northern Chile. Aquaculture International
8, 237–247.
Stotz, W. 2004. Sea urchin fisheries: a Chilean perspective. In Sea Urchins: Fisheries and Ecology, J.M.
Lawrence & O. Guzmán (eds). Lancaster, Pennsylvania: DEStech Publications, 3–17.
337
MARTIN THIEL ET AL.
Stotz, W. & Aburto, J. 2006. Diseño de plan piloto de ordenamiento para pesquerías bentónicas en la IV
Región. Project FIP No. 2003-15, final report. Valparaíso, Chile. Available HTTP: http//www.fip.cl.
Accessed 21 February 2007.
Stotz, W., Aburto, J. & Jaramillo, E. 2004. Ordenamiento de la pesquería de machas de la VIII Región. Project
FIP No. 2003-17, final report, Valparaíso. Available HTTP: http://www.fip.cl. Accessed 21 February 2007.
Stotz, W., Aburto, J. & Lancellotti, D.A. 1999. Estudio de la situación base del área (ESBA) y plan de manejo
y explotación de recursos. Proyecto de manejo y explotación de recursos marinos bentónicos en el
área Tongoy. Unpublished Report, Universidad Católica del Norte, Coquimbo, Chile.
Stotz, W., De Amesti, P., Martínez, D. & Pérez, E.P. 1991b. Lugares de asentamiento y desarrollo de juveniles
tempranos de Concholepas concholepas (Bruguiére, 1789) en el inter y submareal de la IV Región,
Coquimbo. Revista de Biología Marina, Valparaíso (Chile) 26, 339–350.
Stotz, W. & González, S.A. 1997. Abundance, growth, and production of the sea scallop Argopecten purpuratus
(Lamarck 1819): bases for sustainable exploitation of natural scallop beds in north-central Chile.
Stotz, W., González, S.A., Caillaux, L. & Aburto, J. 2003. Quantitative evaluation of the diet and feeding behavior
of the carnivorous gastropod, Concholepas concholepas (Bruguiere, 1789) (Muricidae) in subtidal
habitats in the southeastern Pacific upwelling system. Journal of Shellfish Research 22, 147–164.
Stotz, W., González, S.A. & López, C. 1992. Repoblación experimental del erizo rojo Loxechinus albus Molina
en la costa expuesta del centro-norte de Chile: efectos del erizo negro Tetrapygus niger Molina sobre
el establecimiento y crecimiento de los juveniles. Investigaciones Pesqueras (Chile) 37, 107–117.
Stotz, W., Lancellotti, D.A., Martínez, D., De Amesti, P. & Pérez, E.P. 1991a. Variación temporal del registro
de juveniles recién asentados de Concholepas concholepas (Bruguiére, 1789), en el intermareal rocoso
de Totoralillo, Coquimbo, IV Región. Revista de Biología Marina, Valparaíso (Chile) 26, 351–361.
Stotz, W. & Mendo, J. 2001. Pesquerías, repoblamiento y manejo de bancos naturales, su interacción con la
acuacultura. In Los Moluscos Pectínidos de Iberoamérica: Ciencia y Acuicultura, A.N. Maeda-
Martínez (ed.). Mexico City: Editorial Limusa, 357–374.
Stotz, W. & Pérez, E.P. 1992. Crecimiento y productividad del loco Concholepas concholepas (Bruguiére, 1789)
como estimador de la capacidad de carga en áreas de manejo. Investigaciones Pesqueras (Chile) 37, 13–22.
Strathmann, R.R. 1974. The spread of sibling larvae of sedentary marine invertebrates. American Naturalist
108, 29–44.
Strathmann, R.R. 1990. Why life histories evolve differently in the sea. American Zoologist 30, 197–207.
Strub, P.T. & James, C. 2002. The 1997–1998 oceanic El Niño signal along the southeast and northeast Pacific
boundaries an altimetric view. Progress in Oceanography 54, 439–458.
Strub, P.T., Kosro, P.M. & Huyer, A. 1991. The nature of the cold filaments in the California Current System.
Journal of Geophysical Research-Oceans 96, 14743–14768.
Strub, P.T., Mesías, J.M. & James, C. 1995. Satellite observations of the Perú-Chile counter-current. Geo-
physical Research Letters 22, 211–214.
Strub, P.T., Mesías, J.M., Montecino, V., Rutllant, J. & Salinas, S. 1998. Coastal ocean circulation off western
South America. In The Sea, A.R. Robinson & K.H. Brink (eds). New York: John Wiley and Sons, 273–314.
Stuardo, J. 1964. Distribución de los moluscos marinos litorales en Latinoamérica. Boletín del Instituto de
Biología Marina (Argentina) 7, 79–91.
Stuardo, J. & Valdovinos, C. 1988. A new bathyal Calyptogena from off the coast of central Chile (Bivalvia:
Vesicomyidae). Venus 47, 241–250.
SUBPESCA. 1999a. Cuota global anual de captura 2000 para la pesquería del camarón nailon de la II a la
VIII Región. Informe Técnico (R. Pesq.) No. 63. Valparaíso, Chile.
SUBPESCA. 1999b. Cuota global anual de captura 2000 para la pesquería del langostino amarillo de la III
y IV Región. Informe Técnico (R. Pesq.) No. 62. Valparaíso, Chile.
SUBPESCA. 2005a. Cuota global anual de captura de camarón nailon (Heterocarpus reedi) entre la II y la
VIII Región, año 2006. Informe Técnico de Pesca (R. Pesq.) No. 135. Valparaíso, Chile.
SUBPESCA. 2005b. Cuota global anual langostino amarillo (Cervimunida johni) III y IV Región, año 2006.
Informe Técnico de Pesca (R. Pesq.) No. 134. Valparaíso, Chile.
Sutherland, W.J., Pullin, A.S., Dolman, P.M. & Knight, T.M. 2004. The need for evidence based conservation.
338
Swearer, S.E., Caselle, J.E., Lea, D.W. & Warner, R.R. 1999. Larval retention and recruitment in an island
population of a coral reef fish. Nature 402, 799–802.
Swearer, S.E., Shima, J.S., Hellberg, M.E., Thorrold, S.R., Jones, G.P., Robertson, D.R., Morgan, S.G., Selkoe,
K.A., Ruiz, G.M. & Warner, R.R. 2002. Evidence of self recruitment in demersal marine populations.
Bulletin of Marine Science 70, 251–271.
Takesue, R.K., van Geen, A., Carriquiry, J.D., Ortíz, E., Godínez-Orta, L., Granados, I., Saldívar, M., Ortlieb,
L., Escribano, R., Guzmán, N., Castilla, J.C., Varas, M., Salamanca, M. & Figueroa, C. 2004. Influence
of coastal upwelling and El Niño-Southern Oscillation on nearshore water along Baja California and
Chile: shore-based monitoring during 1997–2000. Journal of Geophysical Research-Oceans 109,
C03009, DOI:10.1029/2003JC001856.
Tala, F. & Edding, M. 2005. Growth and loss of distal tissue in blades of Lessonia nigrescens and Lessonia
trabeculata (Laminariales). Aquatic Botany 82, 39–54.
Tala, F., Edding, M. & Vásquez, J.A. 2004. Aspects of the reproductive phenology of Lessonia trabeculata
(Laminariales: Phaeophyceae) from three populations in northern Chile. New Zealand Journal of
Marine and Freshwater Research 38, 255–266.
Tarazona, J., Arntz, W.E. & Canahuire, E. 1996. Impact of two ‘‘El Niño” events of different intensity on the
hypoxic soft bottom macrobenthos off the central Peruvian coast. Marine Ecology Pubblicazioni Della
Stazione Zoologica Di Napoli I 17, 425–446.
Tarazona, J., Gutiérrez, D., Paredes, C. & Indacochea, A. 2003. Overview and challenges of marine biodiversity
research in Peru. Gayana 67, 206–231.
Tarazona, J., Salzwedel, H. & Arntz, W.E. 1988a. Oscillations of macrobenthos in shallow waters of the
Peruvian central coast induced by El Niño 1982–1983. Journal of Marine Research 46, 593–611.
Tarazona, J, Salzwedel, H. & Arntz, W.E. 1988b. Positive effects of El Niño on macrozoobenthos inhabiting
hypoxic areas of the Peruvian upwelling system. Oecologia 76, 184–190.
Teske, P.R., McQuaid, C.D., Froneman, P.W. & Barker, N.P. 2006. Impacts of marine biogeographic boundaries
on phylogeographic patterns of three South African estuarine crustaceans. Marine Ecology Progress
Series 314, 283–293.
Thiel, M. 2002. The zoogeography of algae-associated peracarids along the Pacific coast of Chile. Journal of
Biogeography 29, 999–1008.
Thiel, M., González, E., Balanda, M.J., Haye, P., Heard, R. & Watling, L. 2004. Diversity of Chilean peracarids
(Crustacea: Malacostraca). In Contributions to the Study of East-Pacific Crustaceans, M.E. Hendrickx
(ed.). Mazatlán, México: Instituto de Ciencias del Mar y Limnología, 177–189.
Thiel, M. & Gutow, L. 2005. The ecology of rafting in the marine environment. II. The rafting organisms and
community. Oceanography and Marine Biology: An Annual Review 43, 279–418.
Thiel, M. & Haye, P. 2006. The ecology of rafting in the marine environment. III. Biogeographical and
evolutionary consequences. Oceanography and Marine Biology: An Annual Review 44, 323–428.
Thiel, M. & Ullrich, N. 2002. Hard rock versus soft bottom: the fauna associated with intertidal mussel beds
on hard bottoms along the coast of Chile, and considerations on the functional role of mussel beds.
Thiel, M. & Vásquez, J.A. 2000. Are kelp holdfasts islands on the ocean floor? Indication for temporarily
closed aggregations of peracarid crustaceans. Hydrobiologia 440, 45–54.
Thomas, A.C. 1999. Seasonal distributions of satellite-measured phytoplankton pigment concentration along
the Chilean coast. Journal of Geophysical Research-Oceans 104, 25,877–25,890.
Thomas, A.C., Blanco, J.L., Carr, M.E., Strub, P.T. & Osses, J. 2001b. Satellite-measured chlorophyll and
temperature variability off northern Chile during the 1996–1998 La Niña and El Niño. Journal of
Geophysical Research-Oceans 106, 899–915.
Thomas, A.C., Carr, M.E. & Strub, P.T. 2001a. Chlorophyll variability in Eastern Boundary Currents. Geo-
physical Research Letters 28, 3421–3424.
Thomas, A.C., Strub, P.T., Carr, M.E. & Weatherbee, R. 2004. Comparisons of chlorophyll variability between
the four major global Eastern Boundary Currents. International Journal of Remote Sensing 25,
1443–1447.
Thorson, G.L. 1950. Reproductive and larval ecology of marine bottom invertebrates. Biological Reviews of
the Cambridge Philosophical Society 25, 1–45.
339
MARTIN THIEL ET AL.
Thurston, M.H., Petrillo, M. & Della Croce, N. 2002. Population structure of the necrophagous amphipod
Eurythenes gryllus (Amphipoda: Gammaridea) from the Atacama Trench (south-east Pacific Ocean).
Tognelli, M.F., Silva-García, C., Labra, F.A., & Marquet, P.A. 2005. Priority areas for the conservation of
coastal marine vertebrates in Chile. Biological Conservation 126, 420–428.
Tokeshi, M. & Romero, L. 1995a. Filling a gap — dynamics of space occupancy on a mussel-dominated
subtropical rocky shore. Marine Ecology Progress Series 119, 167–176.
Tokeshi, M. & Romero, L. 1995b. Quantitative analysis of foraging behavior in a field population of the South
American sun star Heliaster helianthus. Marine Biology 122, 297–303.
Tomanek, L. 2001. The heat-shock response and patterns of vertical zonation in intertidal Tegula congeners.
American Zoologist 41, 1608–1608.
Tomicic, J.J. 1985. Efectos del fenómeno de El Niño 1982–1983 en las comunidades litorales de la península
de Mejillones. Investigaciones Pesqueras (Chile) 32, 209–213.
Torres, R. 1995. Condiciones oceanográficas y baja concentración de clorofila frente a Coquimbo, Chile (Lat.
30°S) durante 1992–1994. M.Sc. thesis, Universidad de Concepción, Concepción, Chile.
Torres, R., Turner, D.R., Rutllant, J. & Lefevre, N. 2003. Continued CO2 outgassing in an upwelling area off
northern Chile during the development phase of El Niño 1997–1998 (July 1997). Journal of Geo-
physical Research-Oceans 108, C10, 3336, DOI:10.1029/2000JC000569.
Tovar, H. & Cabrera, D. 1985. Las aves guaneras y el fenómeno “El Niño”. In “El Niño” Su Impacto en la
Fauna Marina, W. Arntz et al. (eds). Callao, Peru: Instituto del Mar del Perú, 181–186.
Trenberth, K.E. & Hurrel, J.W. 1994. Decadal atmosphere-ocean variations in the Pacific. Climate Dynamics
9, 303–319.
Trillmich, F. 1986. Attendance behavior of Galapagos sea lions. In Fur Seals — Maternal Strategies on Land and
at Sea, R.L. Gentry & G.L. Kooyman (eds). Princeton, New Jersey: Princeton University Press, 196–208.
Trillmich, F. 1990. The behavioral ecology of maternal effort in fur seals and sea lions. Behaviour 114, 3–20.
Troncoso, V.A., Daneri, G., Cuevas, L.A., Jacob, B. & Montero, P. 2003. Bacterial carbon flow in the Humboldt
Current System off Chile. Marine Ecology Progress Series 250, 1–12.
Tyson, R.V. & Pearson, T.H. 1991. Modern and ancient continental shelf anoxia: an overview. In Modern and
Ancient Continental Shelf Anoxia, R.V. Tyson & T.H. Pearson (eds). London: Geological Society
Special Publications, Vol. 58, 1–24.
Ulloa, O. & De Pol, R. 2004. Zonas de mínimo oxígeno. In Biología Marina y Oceanografía: Conceptos y
Procesos, C. Werlinger (ed.). Concepción, Chile: Consejo Nacional del Libro y la Lectura, Universidad
de Concepción, Trama Impresores S.A., 521–532.
Ulloa, O., Escribano, N., Hormazábal, S., Quiñones, R.A., González, R.R. & Ramos, M. 2001. Evolution and
biological effects of the 1997–1998 El Niño in the upwelling ecosystem off northern Chile. Geophys-
ical Research Letters 28, 1591–1594.
Urban, H.J. 1994. Upper temperature tolerance of 10 bivalve species off Perú and Chile related to El Niño.
Uriarte, I., Rupp, G. & Abarca, A. 2001. Spat production of iberoamerican scallops under controlled conditions.
In Los Moluscos Pectínidos de Iberoamérica: Ciencia y Acuicultura, A.N. Maeda-Martínez (ed.).
Mexico City: Editorial Limusa, 147–171.
Uribe, E. & Blanco, J.L. 2001. Carrying capacity of aquatic systems to support intensive scallop aquaculture;
the case of Argopecten purpuratus, Bahía Tongoy, Chile. In Los Moluscos Pectínidos de Iberoamérica:
Ciencia y Acuicultura, A.N. Maeda-Martínez (ed.). Mexico City: Editorial Limusa, 233–248.
Valdés, J., Ortlieb, L. & Sifeddine, A. 2003. Variaciones del sistema de surgencia de Punta Angamos (23°S)
y la zona de mínimo oxígeno durante el pasado reciente. Una aproximación desde el registro sedi-
mentario de la Bahía Mejillones del Sur. Revista Chilena de Historia Natural 76, 347–362.
Valdivia, N., Heidemann, A., Thiel, M., Molis, M. & Wahl, M. 2005. Effects of disturbance on the diversity of
hard-bottom macrobenthic communities on the coast of Chile. Marine Ecology Progress Series 299, 45–54.
Valdivia, N. & Thiel, M. 2006. Effects of point-source nutrient addition and mussel removal on epibiotic
assemblages in Perumytilus purpuratus beds. Journal of Sea Research 56, 271–283.
Valdovinos, C., Navarrete, S.A. & Marquet, P.A. 2003. Mollusk species diversity in the southeastern Pacific:
why are there more species towards the pole? Ecography 26, 139–144.
340
Valle-Levinson, A., Moraga, J., Olivares, J. & Blanco, J.L. 2000. Tidal and residual circulation in a semi-arid
bay: Coquimbo Bay, Chile. Continental Shelf Research 20, 2009–2028.
van Waerebeek, K., Reyes, J. & van Bressem, M. 1998. Sighting of a mother-calf pair of southern right whale
Eubalaena australis in Peruvian waters. Estudios Oceanológicos 17, 105–107.
Vargas, C.A., Araneda, S.E. & Valenzuela, G. 2003. Influence of tidal phase and circulation on larval fish
distribution in a partially mixed estuary, Corral Bay, Chile. Journal of the Marine Biological Associ-
ation of the United Kingdom 83, 217–222.
Vargas, C.A. & Castro, L.R. 2001. Spawning of the chilean hake (Merluccius gayi) in the upwelling system
off Talcahuano in relation to oceanographic features. Scientia Marina 65, 101–110.
Vargas, C.A., Escribano, R. & Poulet, S.A. 2006b. Phytoplankton diversity determines time-windows for
successful zooplankton reproductive pulses. Ecology 87, 2992–2999.
Vargas, C.A. & González, H.E. 2004. Plankton community structure and carbon cycling in a coastal upwelling
system. II. Microheterotrophic pathway. Aquatic Microbial Ecology 34, 165–180.
Vargas, C.A., Manríquez, P.H. & Navarrete, S.A. 2006a. Feeding by larvae of intertidal invertebrates: assessing
their position in pelagic food webs. Ecology 87, 444–457.
Vargas, C.A., Martínez, R., Cuevas, L.A., Pavéz, M., Cartes, C., González, H.E., Escribano, R. & Daneri, G.
2007. The relative importance of microbial and classical food webs in a highly productive coastal
upwelling area. Limnology and Oceanography (in press).
Vargas, C.A., Narváez, D.A., Piñones, A., Navarrete, S.A. & Lagos, N.A. 2006c. River plume dynamic
influences transport of barnacle larvae in the inner shelf of central Chile. Journal Marine Biological
Vargas, C.A., Narváez, D.A., Piñones, A., Venegas, R.M. & Navarrete, S.A. 2004. Internal tidal bore warm
fronts and settlement of invertebrates in central Chile. Estuarine Coastal and Shelf Science 61, 603–612.
Vargas, C.A., Valenzuela, G.S., Núñez, S.P. & Arcos, D.F. 1997. Role of oceanographic and topographic
factors in the retention of hake (Merluccius gayi gayi Guichenot, 1848) larvae in the upwelling system
off central-southern Chile. Archive of Fishery and Marine Research 45, 201–222.
Vásquez, J.A. 1992. Lessonia trabeculata a subtidal bottom kelp in northern Chile: a case study for a structural
and geographical comparison. In Coastal Plant Communities of Latin America, U. Seeliger (ed.). San
Diego, California: Academic Press, 77–89.
Vásquez, J.A. 1993a. Patrones de distribución de poblaciones submareales de Lessonia trabeculata (Laminariales,
Phaeophyta) en el norte de Chile. Facultad de Ciencias del Mar. UCN. Serie Ocasional 2, 187–211.
Vásquez, J.A. 1993b. Abundance, distribution patterns and diets of main herbivorous and carnivorous species
associated to Lessonia trabeculata kelp beds in northern Chile. Facultad de Ciencias del Mar. UCN.
Serie Ocasional 2, 213–229.
Vásquez, J.A. 1995. Ecological effects of brown seaweed harvesting. Botanica Marina 38, 251–257.
Vásquez, J.A. 1999. The effect of harvesting of brown seaweeds: a social, ecological and economical important
resource. World Aquaculture 30, 19–22.
Vásquez, J.A. 2006. Pesca de Investigación: Caracterización de la pesquería de algas pardas de las regiones
I a IV. Temporada 2005–2006. Subsecretaría de Pesca de Chile, Informe Mensual, Coquimbo, Chile.
Vásquez, J.A. & Buschmann, A.H. 1997. Herbivore-kelp interactions in Chilean subtidal communities: a
review. Revista Chilena de Historia Natural 70, 41–52.
Vásquez, J.A., Camus, P.A. & Ojeda, F.P. 1998. Diversidad, estructura y funcionamiento de ecosistemas
rocosos del norte de Chile. Revista Chilena de Historia Natural 71, 479–499.
Vásquez, J.A., Castilla, J.C. & Santelices, B. 1984. Distributional pattern and diets of four species of sea
urchin in a giant kelp forest (Macrocystis pyrifera) of Puerto Toro, Navarino Island, Chile. Marine
Vásquez, J.A., Fonck, E.A. & Vega, J.M.A. 2001a. Diversidad, abundancia y variabilidad temporal de ensam-
bles de macroalga del submareal rocoso del norte de Chile. In Sustentabilidad de la Biodiversidad.
Un Problema Actual: Bases Científico Técnicas, Teorizaciones y Proyecciones, K. Alveal & T. Antezana
(eds). Concepción, Chile: Universidad de Concepción, 351–365.
Vásquez, J.A., Matsuhiro, B., Vega, M.A., Pardo, L.M. & Véliz, D. 2000. The effects of mining pollution on
subtidal habitats of northern Chile. International Journal of Environment and Pollution 13, 453–472.
341
MARTIN THIEL ET AL.
Vásquez, J.A., Pardo, L.M. & Véliz, D. 2001c. Vida bajo las grandes algas. In Sustentabilidad de la Biodi-
versidad. Un Problema Actual: Bases Científico Técnicas, Teorizaciones y Proyecciones, K. Alveal
& T. Antezana (eds). Concepción, Chile: Universidad de Concepción, 351–365.
Vásquez, J.A. & Santelices, B. 1984. Comunidades de macroinvertebrados en discos adhesivos de Lessonia
nigrescens Bory (Phaeophyta) en Chile central. Revista Chilena de Historia Natural 57, 131–154.
Vásquez, J.A. & Santelices, B. 1990. Ecological effects of harvesting Lessonia (Laminariales, Phaeophyta)
in central Chile. Hydrobiologia 204, 41–47.
Vásquez, J.A. & Tala, F. 1995. Repopulation of intertidal areas with Lessonia nigrescens in northern Chile.
Journal of Applied Phycology 7, 347–349.
Vásquez, J.A. & Vega, J.M.A. 2004a. Comunidades marinas costeras del Parque Nacional Bosque Fray Jorge.
In Historia Natural del Parque Fray Jorge, F.A. Squeo et al. (eds). La Serena, Chile: Ediciones
Universidad de La Serena, 235–252.
Vásquez, J.A. & Vega, J.M.A. 2004b. El Niño 1997–1998 en el norte de Chile: efectos en la estructura y en la
organización de comunidades submareales dominadas por algas pardas. In El Niño-La Niña 1997–2000
y sus Efectos en Chile, S. Avaria et al. (eds). Valparaíso, Chile: Comité Oceanográfico Nacional, 119–135.
Vásquez, J.A. & Vega, J.M.A. 2005. Macroinvertebrados asociados a discos de adhesión de algas pardas:
biodiversidad de comunidades discretas como indicadora de perturbaciones locales y de gran escala.
In Biodiversidad Marina: Valoración, Uso y Perspectivas. ¿Hacia dónde va Chile?, E. Figueroa (ed.).
Santiago, Chile: Ediciones Universitaria, 429–450.
Vásquez, J.A., Vega, J.M.A. & Buschmann, A.H. 2006. Long term variability in the structure of kelp com-
munities in northern Chile and the 1997–1998 ENSO. Journal of Applied Phycology 18, 505–519.
Vásquez, J.A., Vega, J.M.A., Matsuhiro, B. & Urzua, C. 1999. The ecological effects of mining discharges
on subtidal habitats dominated by macroalgae in northern Chile: population and community level
studies. Hydrobiologia 399, 217–229.
Vásquez, J.A., Véliz, D. & Pardo, L.M. 2001b. Biodiversidad de macroinvertebrados bajo las grandes algas. In
Sustentabilidad de la Biodiversidad. Un Problema Actual: Bases Científico Técnicas, Teorizaciones y
Proyecciones, K. Alveal & T. Antezana (eds). Concepción, Chile: Universidad de Concepción, 293–308.
Vásquez, J.A., Véliz, D. & Weissner, R. 1996. Análisis malacológico de un yacimiento de la cultura Huente-
lauquén IV Región, Chile. Gayana Oceanológica 4, 109–116.
Vega, J.M.A. 2005. Dinámica de poblaciones de Macrocystis integrifolia en el norte de Chile. M.Sc. thesis,
Universidad Católica del Norte, Coquimbo, Chile.
Vega, J.M.A. & Meneses, I. 2001. Seasonal and spatial monitoring of productivity and of reproduction of
Chondrus canaliculatus (Gigartinales, Rhodophyta) from Chile. Botanica Marina 44, 571–581.
Vega, J.M.A., Vásquez, J.A. & Buschmann, A.H. 2005. Population biology of the subtidal kelps Macrocystis
integrifolia and Lessonia trabeculata (Laminariales, Phaeophyceae) in an upwelling ecosystem of north-
ern Chile: interannual variability and El Niño 1997–1998. Revista Chilena de Historia Natural 78, 33–50.
Velázquez, I., Kaplan, D., Velasco-Hernández, J.X. & Navarrete, S.A. 2005. Multistability in an open recruit-
ment food web model. Applied Mathematics and Computation 163, 275–294.
Véliz, D., Winkler, F.M., Guisado, C. 2003. Developmental and genetic evidence for the existence of three
morphologically cryptic species of Crepidula in northern Chile. Marine Biology 143, 131–142.
Véliz, K., Edding, M., Tala, F. & Gómez, I. 2006. Effects of ultraviolet radiation on different life cycle stages
of the south Pacific kelps, Lessonia nigrescens and Lessonia trabeculata (Laminariales, Phaeo-
phyceae). Marine Biology 149, 1015–1024.
Verheye, H.M., Hutchings, L., Huggett, J.A. & Painting, S.J. 1992. Mesozooplankton dynamics in the Benguela
ecosystem, with emphasis on the herbivorous copepods. South African Journal of Marine Science 12,
561–584.
Vernberg, F.J. 1962. Comparative physiology: latitudinal effects on physiological properties of animal popu-
lations. Annual Review of Physiology 24, 517–546.
Vetter, E.W. & Dayton, P.K. 1998. Macrofaunal communities within and adjacent to a detritus-rich submarine
canyon system. Deep-Sea Research II 45, 25–54.
Vetter, E.W. & Dayton, P.K. 1999. Organic enrichment by macrophyte detritus, and abundance patterns of
megafaunal populations in submarine canyons. Marine Ecology Progress Series 186, 137–148.
342
Vidal, J. 1976. Copépodos calanoideos epipelágicos de la expedición Marchile II. Gayana (Zoología) 15, 1–98.
Villagrán, C. 1995. El cuaternario en Chile: evidencias de cambio climático. In Cambios Cuaternarios en
América del Sur, J. Argollo & P. Mourguiart (eds). La Paz, Bolivia: ORSTOM, 191–214.
Villa-Martínez, R. & Villagrán, C. 1997. Vegetational history of the swamp forests of the central Chilean coast
during the mid and late Holocene. Revista Chilena de Historia Natural 70, 391–401.
Villarroel, J.C., Acuña, E.H. & Andrade, M.J. 2001. Feeding and distribution of the bigeye flounder Hippo-
glossina macrops off northern Chile. Marine and Freshwater Research 52, 833–841.
Villegas, M.J. 2002. Utilización del hábitat por parte de Lontra felina (Molina, 1782) (Carnivora, Mustelidae)
en isla Choros (IV Región de Chile) en relación con la abundancia y distribución de presas. Honors
Villegas, M.J., Aron, A. & Ebensperger, L.A. 2007. The influence of wave exposure on the foraging activity
of the marine otter, Lontra felina (Molina, 1782) (Carnivora: Mustelidae) in northern Chile. Journal
of Ethology DOI 10.1007/s10164-006-0024-x.
Villouta, E. & Santelices, B. 1984. Estructura de la comunidad submareal de Lessonia (Phaeophyta, Lami-
nariales) en Chile norte y central. Revista Chilena de Historia Natural 57, 111–122.
Viviani, C.A. 1979. Ecogeografía del litoral chileno. Studies on Neotropical Fauna and Environment 14, 65–123.
von Brand, E., Merino, G.E., Abarca, A. & Stotz, W. 2006. Scallop fishery and aquaculture in Chile. In
Scallops: Biology, Ecology and Aquaculture, 2nd edition. S.E. Shumway & G.J. Parsons (eds).
Amsterdam: Elsevier Science Publishers, 1293–1314.
Walsh, J.J. 1981. A carbon budget for overfishing off Perú. Nature 290, 300–304.
Wang, H.J., Zhang, R.H., Cole, J. & Chavez, F. 1999. El Niño and the related phenomenon Southern Oscillation
(ENSO): the largest signal in interannual climate variation. Proceedings of the National Academy of
Sciences USA 96, 11,071–11,072.
Wares, J.P., Gaines, S.D. & Cunningham, C.W. 2001. A comparative study of asymmetric migration events
across a marine biogeographic boundary. Evolution 55, 295–306.
Weeks, S.J., Currie, B., Bakun, A. & Peard, K.R. 2004. Hydrogen sulphide eruptions in the Atlantic Ocean
off southern Africa: implications of a new view based on SeaWiFS satellite imagery. Deep-Sea
Research I 51, 153–172.
Weichler, T., Garthe, S., Luna-Jorquera, G. & Moraga, J. 2004. Seabird distribution on the Humboldt Current
in northern Chile in relation to hydrography, productivity, and fisheries. Ices Journal of Marine Science
61, 148–154.
Westermeier, R., Müller, D.G., Gómez, I., Rivera, P. & Wenzel, H. 1994. Population biology of Durvillaea
antarctica and Lessonia nigrescens (Phaeophyta) on the rocky shores of southern Chile. Marine
Westermeier, R., Patino, D., Piel, M.I., Maier, I. & Müller, D.G. 2006. A new approach to kelp mariculture
in Chile: production of free-floating sporophyte seedlings from gametophyte cultures of Lessonia
trabeculata and Macrocystis pyrifera. Aquaculture Research 37, 164–171.
Westermeier, R., Rivera, P.J., Chacana, M. & Gómez, I. 1987. Biological bases for management of Iridaea
laminarioides Bory in southern Chile. Hydrobiologia 151/152, 313–328.
Wiencke, C. & tom Dieck, I. 1990. Temperature requirements for growth and survival of macroalgae from
Antarctica and southern Chile. Marine Ecology Progress Series 59, 157–170.
Wieters, E.A. 2005. Upwelling control of positive interactions over mesoscales: a new link between bottom-
up and top-down processes on rocky shores. Marine Ecology Progress Series 301, 43–54.
Wieters, E.A., Kaplan, D.M., Navarrete, S.A., Sotomayor, A., Largier, J., Nielsen, K.J. & Véliz, F. 2003.
Alongshore and temporal variability in chlorophyll a concentration in Chilean nearshore waters.
Wild, C., Hüttel, M., Klueter, A., Kremb, S., Rasheed, M. & Jorgensen, B. 2004. Coral mucus functions as
an energy carrier and particle trap in the reef ecosystem. Nature 428, 66–70.
Wing, S.R., Botsford, L.W., Largier, J.L. & Morgan, L.E. 1995a. Spatial structure of relaxation events and crab
settlement in the northern California upwelling system. Marine Ecology Progress Series 128, 199–211.
Wing, S.R., Largier, J.L., Botsford, L.W. & Quinn, J.F. 1995b. Settlement and transport of benthic invertebrates
in an intermittent upwelling region. Limnology and Oceanography 40, 316–329.
343
MARTIN THIEL ET AL.
Wishner, K.F., Gowing, M.M. & Gelfman, C. 1998. Mesozooplankton biomass in the upper 1000 m in the
Arabian Sea: overall seasonal and geographic patterns, and relationship to oxygen gradients. Deep-
Sea Research II 45, 2405–2432.
Wishner, K.F., Levin, L., Gowing, M.M. & Mullineaux, L. 1990. Involvement of the oxygen minimum in
benthic zonation on a deep seamount. Nature 346, 57–59.
Woese, C.R. & Fox, G.E. 1977. Phylogenetic structure of the prokaryotic domain: the primary kingdoms.
Proceedings of the National Academy of Sciences USA 74, 5088–5090.
Wolff, M. 1987. Population dynamics of the Peruvian scallop Argopecten purpuratus during the El Niño
phenomenon of 1983. Canadian Journal of Fisheries and Aquatic Science 44, 1684–1691.
Wolff, M. 1988. Spawning and recruitment in the Peruvian scallop Argopecten purpuratus. Marine Ecology
Wolff, M. & Alarcón, E. 1993. Structure of a scallop Argopecten purpuratus (Lamarck, 1819) dominated
subtidal macro-invertebrate assemblage in northern Chile. Journal of Shellfish Research 12, 295–304.
Wolff, M. & Mendo, J. 2000. Management of the Peruvian bay scallop (Argopecten purpuratus) metapopu-
lation with regard to environmental change. Aquatic Conservation-Marine and Freshwater Ecosystems
10, 117–126.
Wood, L.J. 2006. Summary report of the current status of the global Marine Protected Area network, and of
progress monitoring capabilities. Information document provided to the Eighth Ordinaty Meeting of
the Conference of the Parties to the Convention on Biological Diversity. Curitiba, Brazil:
UNEP/CBD/COP/8/INF/4.
Wyrtki, K. 1973. Physical oceanography of the Indian Ocean. In The Biology of the Indian Ocean, B. Zeitzschel
(ed.). Berlin: Springer-Verlag, 18–36.
Yaikin, J., Quiñones, R.A. & González, R.R. 2002. Aerobic respiration rate and anaerobic enzymatic activity
of Petrolisthes laevigatus (Anomura, Porcellanidae) under laboratory conditions. Journal of Crusta-
cean Biology 22, 345–352.
Yannicelli, B. 2005. Distribución y transporte de larvas de crustáceos decápodos en la zona de surgencia
costera de Chile central: interacciones entre el comportamiento, tolerancias fisiológicas y períodos
de liberación. Ph.D. thesis, Universidad de Concepción, Concepción, Chile.
Yannicelli, B., Castro, L.R., Schneider, W. & Sobarzo, M. 2006b. Crustacean larvae distribution in the coastal
upwelling zone off central Chile. Marine Ecology Progress Series 319, 175–189.
Yannicelli, B., Castro, L.R., Valle-Levinson, A., Atkinson, L. & Figueroa, D. 2006a. Vertical distribution of
decapod larvae in the entrance of an equatorward facing bay of central Chile: implications for transport.
Yáñez, E., Barbieri, M.A., Silva, C., Nieto, K. & Espíndola, F. 2001. Climate variability and pelagic fisheries
in northern Chile. Progress in Oceanography 49, 581–596.
Yáñez, E., Catasti, V., Barbieri, M. & Bohm, G. 1996. Relationships between the small pelagic resources
distribution and the sea surface temperatures recorded by NOAA satellites from Chile central zone.
Investigaciones Marinas, Valparaíso 24, 107–122.
Yuras, G., Ulloa, O. & Hormazábal, S. 2005. On the annual cycle of coastal and open ocean satellite chlorophyll
off Chile (18–40°S). Geophysical Research Letters 32, L23604, DOI:10.1029/2005GL023946.
Zacherl, D.C., Manríquez, P.H., Paradis, G., Day, R.W., Castilla, J.C., Warner, R.R., Lea, D.W. & Gaines,
S.D. 2003. Trace elemental fingerprinting of gastropod statoliths to study larval dispersal trajectories.
Zagal, C. & Hermosilla, C. 2001. Guide to Marine Invertebrates of Valdivia. Valdivia: Universidad Austral de Chile.
Zamora, S. & Stotz, W. 1994. Cultivo y producción masiva de juveniles de erizo rojo chileno Loxechinus
albus (Molina, 1782) en laboratorio. Investigaciones Pesqueras 38, 37–54.
Zamorano, J.H., Moreno, C.A. & Duarte, W.E. 1995. Postsettlement mortality in Phragmatopoma virgini (Poly-
chaeta, Sabellariidae) at the Mehuin marine reserve, Chile. Marine Ecology Progress Series 127, 149–155.
Zhang, Y., Wallace, J.M. & Battisti, D.S. 1997. ENSO-like interdecadal variability: 1900–1993. Journal of
Climate 10, 1004–1020.
Zuta, S. & Guillén, O. 1970. Oceanografía da las aguas costeras del Perú. Boletín Instituto del Mar de Perú
2, 157–324.
344

Taylor & Francis
LOSS, STATUS AND TRENDS FOR COASTAL MARINE

HABITATS OF EUROPE
LAURA AIROLDI1 & MICHAEL W. BECK2
1Dipartimento di Biologia Evoluzionistica Sperimentale and
Centro Interdipartimentale di Ricerca per le Scienze Ambientali in Ravenna,

University of Bologna, Via S. Alberto 163, 48100 Ravenna, Italy
E-mail: laura.airoldi@unibo.it
2The Nature Conservancy and Institute of Marine Sciences,
100 Shaffer Road–LML, University of California, Santa Cruz, California 95060, U.S.
Abstract Over the centuries, land reclamation, coastal development, overfishing and pollution
have nearly eliminated European wetlands, seagrass meadows, shellfish beds, biogenic reefs and
other productive and diverse coastal habitats. It is estimated that every day between 1960 and 1995,
a kilometre of European coastline was developed. Most countries have estimated losses of coastal
wetlands and seagrasses exceeding 50% of the original area with peaks above 80% for many regions.
Conspicuous declines, sometimes to virtual local disappearance of kelps and other complex macro-
algae, have been observed in several countries. A few dominant threats have led to these losses
over time. The greatest impacts to wetlands have consistently been land claim and coastal devel-
opment. The greatest impacts to seagrasses and macroalgae are presently associated with degraded
water quality while in the past there have been more effects from destructive fishing and diseases.
Coastal development remains an important threat to seagrasses. For biogenic habitats, such as oyster
reefs and maerls, some of the greatest impacts have been from destructive fishing and overexploi-
tation with additional impacts of disease, particularly to native oysters. Coastal development and
defence have had the greatest known impacts on soft-sediment habitats with a high likelihood that
trawling has affected vast areas. The concept of ‘shifting baselines’, which has been applied mostly
to the inadequate historical perspective of fishery losses, is extremely relevant for habitat loss more
generally. Most habitat loss estimates refer to a relatively short time span primarily within the last
century. However, in some regions, most estuarine and near-shore coastal habitats were already
severely degraded or driven to virtual extinction well before 1900. Native oyster reefs were
ecologically extinct by the 1950s along most European coastlines and in many bays well before
that. These shellfish reefs are among the most endangered coastal habitats, but they receive some
of the least protection. Nowadays less than 15% of the European coastline is considered in ‘good’
condition. Those fragments of native habitats that remain are under continued threat, and their
management is not generally informed by adequate knowledge of their distribution and status.
There are many policies and directives aimed at reducing and reversing these losses but their overall
positive benefits have been low. Further neglecting this long history of habitat loss and transfor-
mation may ultimately compromise the successful management and future sustainability of those
few fragments of native and semi-native coastal habitats that remain in Europe.
345
LAURA AIROLDI & MICHAEL W. BECK
Introduction
Habitat modification, fragmentation and loss are widely considered some of the most serious threats
to diversity globally (Sih et al. 2000). In terrestrial environments, understanding and abating the
effects of habitat loss and fragmentation are a huge focus in science, conservation and management
(Wilcove et al. 1998, Brooks et al. 2002). It has been estimated that, throughout history, humans
have severely modified or exploited to complete loss >70% of natural habitats in the habitable
portion of the planet (Hannah et al. 1994) and that we are still losing somewhere between 0.5%
and 1.5% of wild nature each year (Balmford et al. 2003). Habitat loss is also well recognised as
an important threat in the marine environment (Suchanek 1994, Gray 1997, Wolff 2000) but has
not been as much a focus of science and conservation as in terrestrial environments.
Habitat loss is particularly severe in coastal marine ecosystems, where human activities have
been historically concentrated (Suchanek 1994, Lotze 2004, Knottnerus 2005, Lotze et al. 2006,
Valiela 2006). Coastal zones occupy <15% of the Earth’s land surface, but they accommodate >60%
of the world’s population (EEA 1999a). Globally, the number of people living within 100 km of
the coast increased from roughly 2 billion in 1990 to 2.2 billion in 1995 (Burke et al. 2001), and
the population living on the coast is projected to double in the next 30 yr with an expected 75%
of the world’s population residing in coastal areas by 2025 (EEA 1999a). As human population
has increased in coastal areas, so has the pressure on coastal ecosystems through habitat conversion,
increased pollution, and demand for coastal resources. Coastal systems provide many important
services to humans such as nutrient cycling, food production, provision of habitat/refugia, distur-
bance regulation, natural barriers to erosion, control of water quality, and nursery grounds. Indeed
the global value of services from seagrasses, estuaries and coastal wetlands is estimated to be
10 times higher than that of any terrestrial ecosystems (Costanza et al. 1997).
Recent reviews have examined the extent of habitat loss and fragmentation in tropical environ-
ments across large regions for coral reefs (e.g., Sebens 1994, Spalding et al. 2001, Pandolfi et al.
2003, Wilkinson 2004) and mangroves (e.g., Burke et al. 2001, Valiela et al. 2001, Alongi 2002,
Wilkie & Fortuna 2003). These studies have done much to advance our understanding of the status
and trends of tropical marine ecosystems at multinational and global levels and have been influential
in galvanising support for tropical science, conservation and management.
Our understanding of the status and trends of temperate marine habitats is surprisingly further
behind. Few scientific institutions, organisations or agencies have programmes that focus on tem-
perate marine environments beyond a regional level, and almost no non-governmental organisations
(NGOs) or agencies have multinational or global programmes that focus particularly on temperate
ecosystems such as seagrasses, salt marshes or oyster reefs or the issue of habitat loss. There have
been a few broad reviews of the condition of key temperate habitats (e.g., Kennish 2002, Steneck
et al. 2002, Thompson et al. 2002, Lotze et al. 2006) and some recent exemplary efforts to pull
together global distribution data on seagrasses (Short & Wyllie-Echeverria 1996, Duarte 2002,
Green & Short 2003). Nonetheless, huge gaps still remain in our knowledge of habitat loss on
temperate coasts and estuaries. This gap is particularly disturbing because these coasts contain
some of the most productive, diverse and, at the same time, degraded ecosystems on Earth (Suchanek
1994, Edgar et al. 2000).
In Europe, there has been increasing awareness and concern about the degradation of natural
habitats (e.g., Laffoley 2000). Many European coastal habitats have been lost or severely degraded,
and it is estimated that only a small percentage of the European coastline (<15%) is in ‘good’
condition (EEA 1999a). Unfortunately, there are no comprehensive summaries of the current
distribution and status of marine habitats along European coastlines and even less information is
available about long-term trends of habitat loss or degradation.
346
LOSS, STATUS AND TRENDS FOR COASTAL MARINE HABITATS OF EUROPE
Aim of the review

The aim of the present work is to review up-to-date estimates of large-scale trends in habitat extent,
status and loss along European coastlines. Some of the major causes of these losses and some of
the E.U.-wide policies aimed at slowing these losses are also discussed. Knowledge of the loss of
coastal habitats and the drivers of this change is critical for identifying future directions in the
sustainable conservation and management of Europe’s coastlines (Dayton et al. 1998, Jackson et al.
2001, Lotze 2004).
The review is organised into three major sections: (1) an overview of the historical use of coastal
resources and general impacts to European coastlines; (2) a compilation of data on the current
distribution, historical loss, threats and protection measures of coastal habitats within bays, estuaries
and near-shore shelf environments of Europe; (3) a discussion of gaps in the data, ecological
knowledge and protection measures for these coastal habitats and recommendations for how to
address these gaps.
The information on the coastal habitats of Europe is widely disparate in its availability and
quality. The review of information focuses first and foremost on information that was consistent
and comparable at the Europe-wide level. This information was augmented with data from country-
wide and within-country surveys and occasionally with information from individual bays, estuaries
or sections of country coastlines. The information from these finer levels of resolution mostly
provides key and in-depth examples of coastal change and its causes; it was not possible to collect
this site-specific information for most areas. When possible, information was collected from the
primary literature but much of this information exists in agency reports and online databases and
these were often the most common sources of information.
Definitions
‘Habitat’ and ‘ecosystem’ are terms that have often been used inconsistently and interchangeably
(e.g., Beck et al. 2001). In this review, ‘habitat’ indicates a focus on the predominant features that
create structural complexity in the environment, such as plants (e.g., seagrass meadows, kelp forests),
animals (oyster reefs) or other geological features (e.g., rocky reefs, mudflats). ‘Ecosystem’ indi-
cates a focus on the many other plants, animals, natural processes and services associated with the
predominant features. These definitions are consistent with those commonly used in European
policy (e.g., E.U. Habitats Directive, EC 2003).
‘Habitat loss’ and ‘habitat conversion’ are treated as representations of similar impacts, that is,
a reduction in the distribution of natural habitats. Habitat degradation and fragmentation also
represent serious impacts but they are not often treated as habitat loss because of difficulties in
measurement. Loss clearly occurs when natural habitats such as salt marshes are filled with
sediments and blocked from the sea to form agricultural fields. Habitat conversion often occurs
when more structurally complex natural habitats are converted to less-complex habitats (e.g., oyster
reefs are dredged and mudflats are left). These converted habitats (e.g., dredged mudflats) may still
have some natural value, but they exist for artificial reasons, and less structurally complex habitats
usually have lower diversity and productivity (e.g., Heck & Crowder 1991, Beck et al. 2001). Areas
are rarely converted from less-complex to more complex natural habitats unless there is active
habitat restoration, which is treated as habitat gain. Structurally complex habitats are clearly
becoming rarer across Europe and the world, and that is recognised in their common treatment in
policy, conservation and management.
Habitat degradation, such as the invasion of non-native algae into seagrass meadows or ditching
in marshes, is also a serious issue that has ecosystem implications and often is a precursor to loss
347
Dike
100 m
Creek Ditch Salt marsh
Tidal flat Brushwood groyne
Figure 1 Morphology of a natural salt marsh (left) and an artificial salt marsh (right) functioning as foreland
to protect a dyke in the Wadden Sea. (From Reise 2005. With permission.) Compared with natural ones,
artificial salt marshes are smaller, fragmented, truncated at the landward side and of a simplified structure,
although the halophytic vegetation may be similar.
of natural habitats. Degradation is, however, difficult to measure because it represents a decrease
in condition, not a change in distribution (i.e., habitat loss). Degradation is particularly difficult to
measure at the regional, national and multinational levels considered in this review. Nonetheless it
is clear that present-day salt marshes in Europe, for example, are much different from salt marshes
of the past, not only because they are smaller (habitat loss), but also because they are much less
complex with fewer channels and straighter, less-fractal edges (Figure 1). This degradation results in
much less efficient transfer of nutrients and species at this critical terrestrial/marine interface
(Minello et al. 1994). Habitat fragmentation falls between loss and degradation. Fragmentation
occurs when previously continuous habitats become patchier (e.g., loss of patches of seagrass within
a larger bed). In this review, these changes in the habitat are treated as loss when it can be measured,
which is generally an issue associated with monitoring resolution because many large-scale surveys
and spatial imagery do not capture increases or decreases in patchiness.
General European context

European estuaries and coastal areas have a long history of intense human impacts and urbanisation
and are among the most severely degraded coastal temperate systems worldwide (Lotze et al. 2006).
Europeans have exploited near-shore marine resources since at least the Palaeolithic (e.g., Knott-
nerus 2005), and during the Roman times European coastal landscapes were far from pristine
(Rippon 2000). To improve the habitability of coastal areas and exploit their resources humans
348
B
E
A
H
F
G
C
Figure 2 Native coastal habitats such as oyster reefs (A), salt marshes (B), and seagrass meadows (C) are
being squeezed out of coastal zones by many factors including coastal development and defence (D), diking
and ditching (E), dredging (F), pollution and excess sedimentation (D, E, F), non-native species (G) and
destructive fishing and overfishing (H).
have altered the fluvial sedimentation patterns, controlled the river networks, reclaimed marsh lands
and developed agriculture and fishing (Cencini 1998). Heavy exploitation, modification and dete-
rioration of coastal habitats increased significantly during the Middle Ages, when coastal areas
became more populated and humans started to systematically transform the coastal environment
and commercially exploit its resources (Wolff 1992, Hoffmann 2005), and assumed dramatic
proportions during the nineteenth and twentieth centuries, when uncontrolled coastal development,
industry and tourism destroyed near-shore habitats and assemblages and deeply modified coastal
landscapes and seascapes (Cencini 1998, Reise 2005). This section does not intend to give detailed
information on the history of human exploitation of coastal resources or provide an extensive review
of every type of impact. Rather, it provides baseline information and some key examples of the
major past and present human pressures to European coastlines (Figure 2). This information is
relevant to an understanding of how the concentration of population, settlements and economic
interests in near-shore coastal areas and bays has produced, and still produces, drastic and probably
irreversible changes to native habitats and assemblages (e.g., Lotze et al. 2005). The section also
provides a broad overview of the main E.U. and trans-national agreements and policies that have
been developed to rectify or reduce damage to European natural habitats and associated species.
Threats to European coastlines

The exponential growth of European populations over time (McEvedy & Jones 1978) has driven
the historical intensification and multiplication of human impacts on coastal habitats. Since ancient
epochs, human densities have peaked along European estuaries and coasts, reflecting their impor-
tance for settlement, trade and transport. Many of Europe’s capital cities are on or close to the
coast, and altogether there are 280 coastal cities with populations above 50,000 (Figure 3). In the
1990s, approximately 200 million Europeans lived within 50 km of coastal waters (Stanners &
Bourdeau 1995; see also EEA 1999a, Frid et al. 2003). Today the coasts of many European countries
are among the fastest-growing areas in terms of social and economic development and it is expected
349
−30° −20° −10° 0° 10° 20° 30° 40° 50° 60° Population in coastal
settlements (2001)
Number of inhabitants
50 000–100 000
60°
100 000–500 000
60° 500 000–2.5 million
> 2.5 million
Outside data coverage
50°
50°
40°
40°
Canary Is. −30° Azores Is.

30° 40°
30°
Madeira Is. 0 500 1000 1500 Km

20° 30°
Figure 3 Coastal settlements with more than 50,000 inhabitants along European coasts. (From EEA 2006a.
With permission.)
that European coastal areas will face increasing pressures from population growth (EEA 2005,
2006a). The coasts of the Mediterranean Sea, in particular, have always been among the most
densely populated regions on Earth, with an estimated 5700–6600 people km−1 of coastline in 2000
(UNEP/MAP/PAP 2001). Along Mediterranean coasts, the population increased by 46% between
1980 (84.5 million) and 2000 (123.7 million), and it is projected to nearly double between 2000
and 2025 (UNEP/MAP/PAP 2001).
Increased land use and development of settlements, agriculture, industries, ports, military
installations, mines, power plants and other infrastructures has accompanied population growth in
European coastal areas. Their development has posed and still poses severe threats to coastal areas
(EEA 2006a). Estuarine and coastal landscapes have been deeply modified and transformed in a
process that in some regions, such as the western Netherlands, dates as far back as late prehistoric
periods, when the first attempts were made to control the flow of water through the construction
of dams and sluices (Rippon 2000). During the Roman times, reclamation of coastal marshes
became intensive in some regions (e.g., the Severn Estuary; Rippon 1997), and after the tenth to
twelfth centuries, large-scale transformations and reclamations took place systematically around
Europe (Wolff 1992, Cencini 1998, Rippon 2000, Reise 2005). In the Wadden Sea region, about
350
15,000 km2 of wetland, lagoons, coastal lakes and tidal flats have been embanked, drained and
converted into arable land and pasture over the centuries (Figure 4; see also Wolff 1992, 1997). In
the United Kingdom land reclamation has affected at least 85% of the estuaries since Roman times,
with losses of intertidal areas ranging between 25 and up to >80% (Davidson et al. 1991); such
widespread claim of estuarine land is continuing at rates of 0.2–0.7% yr−1 and affects also estuaries
of recognised international wildlife importance included in the Ramsar/Special Protection Area
(SPA) network.
Data from the CORINE project indicate that 22,000 km2 of the coastal zone in Europe are
covered in concrete or asphalt (EEA 2005), and that artificial surfaces increased by almost 1900 km2
between 1990 and 2000 alone (EEA 2006a). The greatest urban developments occur along the
Euro-Mediterranean coasts. At present about two thirds of the Mediterranean coastline is urbanised,
with this fraction exceeding 75% in the regions with the most developed industries
(UNEP/MAP/PAP 2001). More than 50% of the Mediterranean coasts are dominated by concrete
with >1500 km of artificial coasts, of which about 1250 km are developed for harbours and ports
(EEA 1999c). Growth of cities (particularly tourist developments) and development of industry in
some regions (including the French Riviera, Athens, Barcelona, Marseille, Naples, the north Adriatic
shorelines) have taken up to 90% of the coastline (Jeftic et al. 1990, Meinesz et al. 1991, Cencini
1998). In Italy, a survey carried out by World Wildlife Fund (WWF) showed that, in 1996, 42.6%
of the entire Italian coast was subject to intensive human occupation (areas completely occupied
by built-up centres and infrastructures), 13% had extensive occupation (free zones occupied only
by extensive building and infrastructures) and only 29% was free from buildings and infrastructures
(EEA 1999c). Coastal zone urbanisation will further increase in the near future, with projected
increases of 10–20% for most Mediterranean countries (EEA 2006a).
Severe decreases of water quality have generally followed population growth with organic
pollution as a major driving factor (Jansson & Dahlberg 1999, Diaz 2001, van Beusekom 2005).
Excessive nutrient enrichment has been a problem in European waters historically (Islam & Tanaka
2004). Hoffman (2005) reports that archaeological signs of eutrophication from dense, mainly urban
populations were detected on the Bodensee shore at Konstanz (Germany) in late-mediaeval times,
and that in 1415 a royal ordinance tried to mitigate the low water quality of the Seine below Paris.
Nutrient loads started to rise probably around 1700–1800, increased significantly in the early 1900s
and steeply accelerated after the 1950s (Lotze et al. 2006). It is estimated that in the Baltic and
North Sea regions nitrogen (N) and phosphorus (P) loads from land and atmosphere have increased
about 2–4 and 4–8 times, respectively, since the 1940s (Nehring 1992, EEA 2001, Karlson et al.
2002). Historical reconstructions of the preindustrial trophic status in the Wadden Sea suggest about
5-fold greater organic matter turnovers nowadays compared with preindustrial conditions (van
Beusekom 2005). The historical development in nutrient loads to the Mediterranean and Black
Seas is unknown, but is probably of the same magnitude (UNEP/FAO/WHO 1996, EEA 1999c).
For example, in the north Adriatic Sea nutrient load has been increasing since at least 1900 and it
markedly intensified after 1930 (Barmawidjaja et al. 1995, Sangiorgi & Donders 2004), with a
doubling of nutrient loads in the Po river between 1968 and 1980 (Marchetti et al. 1989). In the
Black Sea, concentrations of nitrate have increased 5 times and phosphate 20 times from the 1960s
to 1980s (Gomoiu 1992).
The increased eutrophication has, as a secondary effect, led to increased oxygen consumption
on the sea bed and expansion of areas with hypoxia and anoxia (Diaz 2001, Karlson et al. 2002).
In the Black Sea up to 90% of the waters are anoxic. The Kattegat has been affected by seasonal
hypoxia since the beginning of the 1980s, which has followed a more than 3-fold increase in N
input in the 1960s and 1970s (Rosenberg et al. 1990). Similarly, in the north Adriatic Sea the first
signs of hypoxia started around 1960 and developed into severe anoxic events over the past 20 yr
(Barmawidjaja et al. 1995, Diaz 2001). Since the middle of the 1980s the phosphorus load has
351
Wieding-
harde
co.1436 56
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Risum-
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üll
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Hingstness Südhörn
Wieding
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6
Figure 4 Maps of about 20 km of coasts in Nordfriesland circa 1500 (top) and in 1965 (bottom). (From Reise
2005. With permission.) Shown is the massive loss of coastal habitats due to land claim. In 1500 Dagebüll
and Fahretoft islands were surrounded by low summer dykes. All the area was subsequently embanked (years
of progressive diking are indicated in the 1965 map), and tidal flats and salt marshes were drained and converted
to arable land and pasture. Pleistocene elevations are hatched, salt marshes stippled, tidal flats dotted, former
creeks narrowly dotted and arrows point to sites of shore erosion.
352
generally levelled off or declined locally. In some areas such as the North Sea there have been
declines in P up to 50% due to improved sewage treatment, reduced industrial discharges and a
change to phosphorus-free detergents (Frid et al. 2003). However, there do not yet seem to be
discernible European-scale reduction of nitrogen inputs, marine eutrophication or extent of anoxic
areas (Karlson et al. 2002).
Increased loads of sediments have followed changes in land use both inland and along the
coasts of Europe, but long-term data on water turbidity and sediment load are limited even at local
scales (Lumb 1990). The greatest impacts were felt when forests were extensively cleared for
timber, agriculture or urban developments, which together with interferences in the natural course
of rivers caused dramatic acceleration of natural soil erosion (Airoldi 2003). In Europe clearing of
forested catchments for agriculture commenced several thousand years ago (e.g., see the historical
reconstruction by Cencini 1998). Episodes of accelerated erosion following phases of expansion
of arable lands were common during mediaeval times (Hoffmann 2005) and became particularly
severe during the nineteenth century (Pukaric & Jorissen 1990, Barmawidjaja et al. 1995).
Chemical pollution has also affected European estuaries and coastal waters since ancient times
(Islam & Tanaka 2004), particularly in the Mediterranean Sea, where overall 101 pollution ‘hot
spots’ have been identified, generally located in semi-enclosed gulfs and bays near important
harbours, big cities and industrial areas (UNEP/MAP/PAP 2001, EEA 2006b). Pollution from
shipping, oil spill traffic, drilling activities and related accidents is particularly severe in Europe
(EEA 1998, 1999c, 2006a, Thompson et al. 2002). Some of the busiest shipping lanes in the world
are found in the Baltic Sea, North Sea and Mediterranean Sea (Frid et al. 2003), and about 22%
of the total world petroleum traffic passes through the Mediterranean Sea (Jeftic et al. 1990). Marine
pollution has become a major concern in Europe, and many E.U., trans-national and national
initiatives (see next section, p. 356) have helped to control the disposal of urban and industrial
pollutants in coastal areas. Even so, there are still large pollution loads, and long-term contamination
of sediments is a major problem.
Marine food resources have been used by Europeans since prehistory. At some heavily populated
localities, particularly along the Mediterranean shores, the most valued species had severely
declined in abundance and size by the end of the Roman era (Hoffmann 2005) with detectable
effects on coastal systems (Sala 2004). Exploitation increased during late-mediaeval times, when
fisheries became subject to market exploitation, and in subsequent centuries growing food demand
and technological progress led to almost unrestricted overexploitation of coastal resources (Hoff-
mann 2005, Wolff 2005, Lotze et al. 2006). The total fish landings in European sea regions peaked
at 12 million t in 1997, but have decreased since in both quantity and quality, down to 7.6 million t
in 2002 (EEA 2006a).
Disruptive fishing techniques are considered among the major causes of physical destruction
of marine coastal habitats at global scales (Watling & Norse 1998, Turner et al. 1999, Thrush &
Dayton 2002). In Europe, bottom trawls, bivalve dredging, pneumatic hammering of date mussels,
explosives and other disruptive fishing techniques have a long history of use, mainly in estuaries, bays
and continental shelf waters (Fanelli et al. 1994, Bavestrello et al. 1997, Lindeboom & de Groot
1998, Cicogna et al. 1999, EEA 1999c, Johnson 2002, Hall-Spencer et al. 2003, Tudela 2004). In
Britain, concern about the adverse effects of fishing on marine habitats and wildlife populations
dates back to the fourteenth century when it was noted in a petition presented to Parliament in the
year 1376–1377 (quoted in Hore & Jex 1880) that “the hard and long iron of the said ‘wondyrchoun’,
[an oyster dredge] … destroys the spawn and brood of the fish beneath the said water, and also
destroys the spat of oysters, muscles [sic], and other fish by which large fish are accustomed to
live and be supported”. The use of trawls and other mobile fishing gears accelerated sharply with
the introduction of diesel engines in the 1920s (Watling & Norse 1998). The sea bed in Europe
353
has been trawled to a depth of over 1000 m since the 1970s, affecting extensive areas of benthic
habitats.
Aquaculture, which can have some benefits, has had increasingly adverse effects on coastal
habitats. World aquaculture production has increased by >300% since 1984, with growth of about
10% a year in the 1990s, making it the fastest-growing food production activity (Mock et al. 1998).
In Europe the culture of fish, shrimp, shellfish and seaweeds has been used as an alternative source
of marine food at least since Roman times (Hoffmann 2005), and in regions such as the Po delta area
salt marshes were transformed centuries ago into artificial fishing lagoons (Cencini 1998). Unprec-
edented growth in production has occurred in the last decades (Váradi 2001, EEA 2006a), with
significant impacts on bottom habitats and assemblages (e.g., Holmer et al. 2001, EEA 2006b). In
1998, total marine aquaculture production in Europe was >1.3 million t, with most production
concentrated in Norway, France, Spain and Italy (Váradi 2001). In the Mediterranean region, marine
aquaculture production has increased from 19,997 t in 1970 to 339,185 t in 2002 (EEA 2006b),
and the total production of salmon in fish farms (mainly in Norway and Scotland) has increased
from 70,000 t in 1990 to 148,000 t in 1996 (EEA 2002) up to 540,000 t in Norway alone in 2003
(EEA 2006a).
More recent pressure and threats to European coastlines are from tourism and development of
recreational infrastructures, particularly in the Mediterranean region. Before the 1930s, tourism
was a relatively minor phenomenon, although it did lead to the beginning of urbanisation in seaside
areas (EEA 1999c). From the 1930s onward and especially after World War II, mass tourism started
to grow, and the phenomenon was amplified by the development of transport facilities (e.g., Cencini
1998). Nowadays, the Mediterranean coast is the world’s leading holiday destination, accounting
for 30% of the world’s tourism, and in some countries coastal tourism represents up to 90% of all
tourism. In 1990 alone, 135 million vacationers flocked to the Mediterranean coast, and by 2025
the annual crowd is projected to increase to 235–350 million tourists (EEA 1999c). Effects on
coastal habitats have been devastating. In Spain, tourist developments occupy 42% of the entire
coast (Jeftic et al. 1990), with peaks in areas such as the Catalonia coast, where tourist developments
make up 337 km of the total 580 km. Similarly, buildings, roads, bathing establishments and other
recreational facilities located directly over the beaches and sand dunes almost entirely occupy the
Italian coast of the north Adriatic Sea (Cencini 1998). The demand for marinas and yacht harbours
has been growing all over the Mediterranean coasts, with an estimated growth for France of
1.5–2.6% yr−1 (EEA 2006a).
Increased coastal erosion and flooding (often indirectly related to human activities) also pose
serious threats to European coastlines (EC 2004). A recent inventory of coastal evolution in Europe
undertaken within the CORINE programme showed 55% of the coastline to be stable, 19% to
be suffering from erosion problems and 8% to be depositional (Stanners & Bourdeau 1995). Some
coastal regions are also gradually subsiding (Bondesan et al. 1995, EEA 2006a), with subsidence
sometimes enhanced by groundwater or petrochemical extraction (Bird 1993), while land lift up
to 9 mm yr−1 is occurring in areas of the Baltic Sea (HELCOM 1998).
Erosion mitigation schemes have been put in place to respond to the problem of coastal erosion,
which affected about 7600 km of coasts in 2001 (EC 2004). Defence measures include a variety
of hard defence structures (e.g., breakwaters, groynes, seawalls, dykes or other rock-armoured
structures), which have proliferated in the second half of the twentieth century, leading to severe
hardening of coastal areas and changes in sediment structure (Airoldi et al. 2005). In the north
Adriatic Sea, >190 km of artificial structures, mainly groynes and breakwaters, seawalls and jetties
(Figure 5), have been built along 300 km of naturally low sedimentary shores (Bondesan et al.
1995, Cencini 1998). This hardening has caused severe losses and alterations of shallow sedimentary
habitats (e.g., Martin et al. 2005) and has introduced new artificial habitats, with dramatic effects
on native habitats and assemblages (Bacchiocchi & Airoldi 2003, Bulleri & Airoldi 2005). Similar
354
Figure 5 Coastal defence structures along the highly urbanised Italian shores of the north Adriatic Sea. (Photo
by Giorgio Benelli. With permission.)
examples occur in many other European coasts (e.g., Davidson et al. 1991, Anthony 1994, Reise
2005), presumably affecting an overlooked enormous amount of benthic habitats.
Overall >15,000 km of coasts in Europe are now actively retreating, some of them in spite of
coastal protection works (2900 km), and another 4700 km are artificially stabilised (EC 2004).
Globally the problem of erosion and flooding is becoming much more serious because of rising
sea levels and an increased storm frequency as a result of global climate change (Bray & Hooke
1997, Valiela 2006). During the past century, the mean global sea level has risen between 10 and
25 cm (Burke et al. 2001). The Intergovernmental Panel on Climate Change (IPCC) Working Group
355
has projected a global sea-level rise of 15–95 cm by the year 2100. The recession of coastlines is
expected to continue even in the absence of new human activities (Bondesan et al. 1995).
Approximately 450–600 non-indigenous marine species have been added to European coastal
fauna and flora, often facilitated by human-mediated processes such as shipping, aquaculture and
aquaria (Reise et al. 2006 and references therein). Some introductions occurred hundreds of years
ago, as is the case of the sand-gaper, Mya arenaria, which was probably transported as food from
North America to Europe by the Vikings (Petersen et al. 1992). Rates of introduction rose dramat-
ically in the past century, probably in relation to increased shipping and aquaculture. In the
Mediterranean Sea, the number of introduced species has nearly doubled every 20 yr since the
beginning of the twentieth century (Boudouresque & Verlaque 2002). The number of introduced
species is often greatest in estuaries, lagoons, embayments, closed seas, canals and harbours,
probably because of low species richness combined with strong anthropogenic change (Occhipinti-
Ambrogi 2001, Reise et al. 2006). They have profoundly altered European coastal ecosystems and
are displacing some native species. One notorious example is the invasion of Caulerpa taxifolia
along the coastlines of Spain, France, Monaco, Italy, Croatia and Tunisia (Meinesz et al. 2001)
although there is debate about the area of benthos affected (Jaubert et al. 2003). Nevertheless, these
invasions do not seem to have caused large-scale extinctions in recipient biota and losses of native
coastal habitats (Wolff 2000).
E.U. coastal policies and trans-national agreements

The European Union has been involved in efforts to protect the European natural heritage for the
past 30 yr (Table 1). At the international level, the European Union has signed a number of important
conventions aimed at nature protection, including the Ramsar Convention on the Conservation of
Wetlands, the Bonn Convention on Migratory Species, and the Rio Convention on Biological
Diversity (CBD), and shares the international commitment of the World Summit for Sustainable
Development to establish a globally representative system of marine and coastal protected areas
by 2012 (Kelleher et al. 1995, Green & Paine 1997).
At the European level, the Bern Convention has led the development of policy and action in
nature conservation in Europe. It lists protected species, including a number of marine plants and
invertebrates, and requires its parties to prevent the disappearance of endangered natural habitats.
The Sixth Environmental Action Plan, setting the European Union’s environmental policy agenda
until 2012, highlighted nature and biodiversity as a top priority, and the E.U. leaders in Gothenburg
in 2001 launched the E.U. Sustainable Development Strategy to halt the loss of biodiversity in the
European Union by 2010. The European Union has adopted a Biodiversity Strategy and Action
Plans (currently under review), and Member States have developed — or are developing — their
own national strategies and action plans (e.g., U.K. Biodiversity Group 1999).
Other policies, in particular the Birds Directive and the Habitats Directive, have been promoted
to rectify or reduce damage to European natural habitats and associated species. Following the
criteria set out in the directives, each Member State must draw up a list of sites hosting wild fauna
and natural habitats and put in place a special management plan to protect them, combining long-
term preservation with economic and social activities, as part of a sustainable development strategy.
The final aims are the creation of a European Ecological Network of Special Protection Areas
(SPAs) and Special Areas of Conservation (SACs), called NATURA 2000, and the integration of
nature protection into other E.U. policy areas, such as agriculture, fishery, industry, regional
development and transport.
Indirect protection to a variety of habitats also comes from a number of E.U. Directives that
regulate water quality, including the Dangerous Substances, Shellfish Waters, Integrated Pollution
356
Table 1 Summary of main protection initiatives adopted by the European Union and State
Members that directly or indirectly address issues related to the protection of European marine
coastal habitats and associated assemblages
Initiative Description Web site
Ramsar Ramsar Convention on Wetlands, Ramsar (1971). www.ramsar.org
Convention Provides the framework for the conservation and wise
use of wetlands of international importance especially
as waterfowl habitat. Includes salt marshes and some
lagoon systems and marine waters to a depth of 6 m.
Bonn Convention on the Conservation of Migratory Species of www.cms.int
Convention Wild Animals, Bonn (1979). Intergovernmental treaty,
aiming to conserve terrestrial, marine and avian
migratory species throughout their range.
Rio Convention Convention on Biological Diversity, Rio de Janeiro www.biodiv.org/convention/default.shtml
(1992). Provides legal framework for biodiversity
conservation and sustainable development. The Jakarta
Mandate (1995) leads activity in marine biodiversity
management and conservation.
Bern Convention Convention on the Conservation of European Wildlife www.coe.int/T/E/Cultural_Co-operation/
and Natural Habitats, Bern (1979). Aims at preserving Environment/Nature_and_biological_
wild flora and fauna (including some marine species) diversity/Nature_protection/
and their natural habitats through national programmes
using the co-operation between European States.
ICES Convention for the International Council for the www.ices.dk/aboutus/convention.asp
Convention Exploration of the Sea, Copenhagen (1964). Co-
ordinates and promotes marine research in the North
Atlantic, including the Baltic Sea and North Sea, and
the Common Fisheries Policy on the protection of the
marine environment and the regulation of fisheries.
OSPAR Convention for the Protection of the Marine Environment www.ospar.org
Convention of the northeast Atlantic, Paris (1992). Merged the 1972
Oslo Convention on dumping waste at sea and the 1974
Paris Convention on land-based sources of marine
pollution. It guides the protection of the marine
environment of the northeast Atlantic and the
identification of priority habitats and species.
North Sea Six declarations produced at as many International www.sweden.gov.se/sb/d/6363/a/57475;
Conference Conferences on the Protection of the North Sea (first in jsessionid=abPgelqjfxJ8
Declarations Bremen, 1984). Political commitments to the protection
of the North Sea environment, addressing, e.g., species
and habitats issues, pollution and fisheries.
Helsinki Convention on the Protection of the Marine Environment www.helcom.fi
Convention of the Baltic Sea Area, Helsinki (1992). Guides the
protection of the marine environment of the Baltic Sea
from pollution and the identification of priority habitats
and species for protection.
Trilateral Joint Declaration of The Netherlands, Denmark and www.waddensea-secretariat.org
Wadden Sea Germany, Copenhagen (1982). Aimed at co-ordinating
Cooperation the protection of the Wadden Sea National Park. In
1997, a Trilateral Wadden Sea Plan was adopted.
357
Table 1 (continued) Summary of main protection initiatives adopted by the European Union
and State Members that directly or indirectly address issues related to the protection of European
marine coastal habitats and associated assemblages
Initiative Description Web site
Barcelona Amended in 1995 as the Convention for the Protection www.unepmap.org
Convention/ of the Marine Environment and the Coastal Region of
Mediterranean the Mediterranean, Barcelona (1976). Provides legal
Action Plan framework to MAP (1975), under UNEP Regional Seas
(MAP) Programme. Aims to control human impacts (e.g.,
marine pollution, tourism) and protect the marine and
coastal Mediterranean environments.
Bucharest Convention on the Protection of the Black Sea against www.blacksea-commission.org
Convention Pollution, Bucharest (1992). Aims to control and
prevent pollution and preserve biodiversity in the Black
Sea.
Birds Directive Council Directive on the Conservation of Wild Birds. www.ec.europa.eu/environment/nature/
(79/409/EEC) Identifies 194 endangered species and subspecies of nature_conservation/eu_nature_legislation/
birds for which the E.U. Member States are required to birds_directive/index_en.htm
designate Special Protection Areas (SPAs). Over 4000
SPAs have been designated to date, covering 8% of E.U.
territory.
Habitats Council Directive on the Conservation of Natural www.ec.europa.eu/environment/nature/
Directive Habitats and of Wild Fauna and Flora. Aims to protect nature_conservation/eu_nature_legislation/
(92/43/EEC) wildlife species and habitats which have conservation habitats_directive/index_en.htm
that requires the designation of Special Areas of
Conservation (SACs). These sites, together with the
SPAs of the Birds Directive, make up the NATURA
2000 network, currently covering about 15% of E.U.
coasts. Marine habitats broadly defined, and few marine
species listed.
Shellfish Waters Council Directive on the Quality Required of Shellfish www.europa.eu.int/eur-lex/en/consleg/pdf/
Directive Waters. Aims to ensure a suitable environment for 1979/en_1979L0923_do_001.pdf
(79/923/EEC) shellfish harvest. Member States are required to
designate coastal and brackish waters that need
improvement to support shellfish fisheries.
Water Integrates and updates existing E.U. water legislations www.europa.eu.int/comm/environment/
Framework (e.g., Discharges of Dangerous Substances, Urban water
Directive Waste Water Treatment, Nitrates Directive) and
(2000/60/EC) provides for water management. Complemented by the
recently revised Bathing Water Directive (2006/7/EC).
Marine Strategy The proposed directive aims to define common objectives www.ec.europa.eu/environment/water/
Directive and principles at E.U. level to achieve good marine.htm
environmental status of the European marine
environments by 2021. It will establish European
Marine Regions as management units for
implementation.
Note: All web sites last accessed 8 August 2006.
358
Control, Urban Waste Waters, Bathing Waters and the Water Framework (Table 1). These commit-
ments provide for the regulation of discharges to the sea and have set targets and quality standards
covering many metals, pesticides and other toxic substances.
In addition to E.U. initiatives, a number of trans-national agreements have been developed to
address some conservation issues within the main European seas, including the OSPAR Convention,
the North Sea Conference Declarations, the Helsinki Convention, the Trilateral Cooperation on the
Protection of the Wadden Sea, the Mediterranean Action Plan and the Black Sea Environmental
Programme (Table 1). These programmes generally address water quality and fishery concerns and
are not specifically focused on habitat loss and protection, although initiatives have also included
commitments toward establishing an integrated network of Marine Protected Areas (MPAs). A more
focused initiative for the Atlantic Ocean and Baltic Sea is the commitment of the Joint Ministerial
Meeting of the Helsinki and OSPAR Commissions to complete by 2010 a joint network of MPAs
that, together with the NATURA 2000 network, would be ecologically coherent.
In recent years there has been increasing awareness that past efforts to protect European marine
coastal habitats and associated species have been marginal relative to terrestrial environments and
that there is limited co-ordination of national and transnational initiatives at a European level. The
global MPA database indicates that there are 1129 MPAs in Europe (Figure 6) covering 236,000 km2
(MPA Global 2006). Most of these MPAs are small, and they often lack adequate political and
0 250 500 1,000

Km
Figure 6 Marine Protected Areas in Europe (UNEP/WCMC 2006; data extracted August 2006). These MPAs
include, for example, nature reserves, national parks, habitat/species management areas, RAMSAR Wetlands
of International Importance and World Heritage Sites. The map should be considered indicative of general
distribution not areal extent of MPAs.
359
financial support and effective enforcement (an extensive review of MPAs in Europe is offered in
Kelleher et al. 1995, and specifically for the Mediterranean Sea in Badalamenti et al. 2000). The
application of the Birds and Habitats Directives to the marine environment is also presenting major
challenges, with significant delays in the selection and designation of the marine sites of the
NATURA 2000 network. In response the European Union has set the protection of the marine
environment as a major priority and has launched the Marine Strategy Directive, specifically aimed
at protecting and conserving marine ecosystems and promoting the sustainable use of marine
resources through the development of an integrated, coherent policy for the marine environment.
A 2004 conference in Malahide (Ireland) has also set the necessary steps to complete the selection
of the marine NATURA 2000 sites by the end of 2008.
Coastal habitats in Europe

Despite its relatively small geographic size, Europe has a very long coastline, approximating
325,892 km, including islands (Pruett & Cimino 2000). It comprises the main marine regions of
the northeast Atlantic, part of the Arctic, the Baltic Sea, the North Sea, the Mediterranean Sea and
the Black Sea (Frid et al. 2003, EEA 2006a). It includes primarily temperate environments as well
as some Arctic and subtropical climate environments and covers a variety of geomorphological
features (EEA 2002).
There are a large variety of extremely productive, diverse and valuable natural coastal habitats,
including sea-bed communities of macroalgae and seagrasses, tidal mudflats, salt marshes and
biogenic reefs (Stanners & Bourdeau 1995). However, there does not seem to be any comprehensive
summary of the current distribution and status of native habitats along European coastlines as a whole.
Databases are available or are being prepared for some habitats (e.g., wetlands (Nivet & Frazier
2004) and seagrasses (Green & Short 2003)), countries (e.g., the United Kingdom; Hiscock &
Tyler-Walters 2006), and regions (e.g., northwest Europe, the OSPAR regions and the Baltic Sea;
ICES 2006a). However, for most European coasts information, if any, is scattered, fragmented and
not easily accessible. Even less information is available about long-term trends of habitat loss or
degradation. With some notable exceptions (e.g., the Wadden Sea; Lotze & Reise 2005), there is
little comprehensive historical information on coastal habitats prior to about 1900 (e.g., Hiscock
& Kimmance 2003).
In this section published information, from both the scientific and grey (reports, web sites)
literature is critically documented for major coastal habitats. The aim was not to reconstruct trends
from local historical sources, maps or databases but to cover the most pertinent literature that
reported data and information at regional, national, trans-national or European levels. The data
summarised in this section are of variable quality and it should not be inferred that they provide a
complete picture of the status of European coastal habitats.
Coastal wetlands and salt marshes

Current distribution and status
Much of the European coastline consists of a chain of extensive estuaries, lagoons and intertidal
bays interspersed through stretches of rocky shore and sandy beaches. These coastal wetlands
represent some of the most productive and biologically diverse components of near-shore ecosys-
tems (Dugan 1993, Keddy 2000). They provide nursery grounds for commercially important fishes,
habitat for shellfish, birds and a variety of biota and play a fundamental role in flood control,
nutrient cycling and sediment dynamics. These coastal wetlands are patchworks of sand, mud flats
360
and salt marshes. Salt marshes, with their vegetated complex surfaces, form one of the most
important components of these wetlands (Adam 1990).
Coastal salt marshes are distinctive habitats that can be delineated relatively easily on remotely
sensed images (e.g., Ekebom & Erkkilä 2003). In Europe, they have been the target of studies for
a long time (see, among others, Dijkema 1984, Allen & Pye 1992, Jones & Hughes 1993, Rippon
2000, Adam 2002). However, quantitative information on their distribution and status is limited
even at local scales. Information is often available for the more general category of ‘coastal
wetlands’ and here the focus is mainly on this broader habitat type. Accounts exist at regional and
local scales but most European countries lack comprehensive inventories of the extent and status
of coastal wetlands, with numerous countries lacking almost any organised information on these
conspicuous habitats (Table 2). One of the greatest difficulties in completing such inventories is in
identifying intact wetlands, that is those wetlands that are not so severely transformed or deteriorated
as to be functionally extinct (Allen 2000, Nivet & Frazier 2004). In the Severn Estuary, England,
for example, there are a total of about 14 km2 of intact marshes and about 840 km2 of enclosed
marshes, and this ratio of intact to enclosed coastal marshes may be common across Ireland, France,
the Netherlands, northwest Germany and Denmark (Allen 2000). Other difficulties in estimating
coverage include incompatible information (e.g., inconsistent classifications and methodologies)
and the lack of co-ordination between different studies or for different wetland types (Nivet &
Frazier 2004).
A European-scale review of the current distribution and coverage of coastal wetlands has been
recently completed by Nivet & Frazier (2004) that integrates and updates previous inventories by
Jones & Hughes (1993) and Finlayson & Spiers (1999). According to this inventory, the total cover
of marine/coastal wetlands in Europe is around 51,910 km2, and detailed information for individual
countries is summarised in Table 2. An inventory of the distribution of European wetlands, including
coastal wetlands, is also available in the CORINE database (EEA 1999b). A broad map of the
distribution of salt marshes in Europe is given in Figure 7.
There is little comprehensive information on the status of coastal wetlands and salt marshes in
Europe but there are clear indications that the historical concentration of human activities in
European coastal wetlands has deeply modified their structure and function (Dijkema 1984, U.K.
Biodiversity Group 1999, Allen & Pye 1992). Adam (2002) points out that nowadays minerogenic
sedimentation prevails over autogenic (organic matter) sedimentation in the majority of European
marshes. Most wetlands have deeply altered flow regimes (e.g., Cencini 1998, Reise 2005) with
associated important effects on sediment dynamics as well as nutrient and salinity regimes (e.g.,
Allen 2003) and often are heavily polluted (e.g., Trombini et al. 2003). Their vegetation composition
is the product of centuries of use and management (e.g., Wolff 2000) and their fauna and flora have
been deeply transformed by introduced species (Reise et al. 2006).
Historical losses and causes
Coastal wetlands have suffered some of the most serious habitat loss rates (Dugan 1993, Suchanek
1994, Rippon 2000, Valiela 2006) and some estimates suggest that over time temperate estuaries
and coastal areas may have lost approximately 67% of the wetlands that existed (Lotze et al. 2006).
Even when wetlands have not been completely lost, significant degradation of their environment
has often occurred, impairing their functions (Dugan 1993, Wolff 2000).
Exploitation of coastal wetlands and salt marshes in Europe dates back to at least the Neolithic,
when salt marshes were used for salt production (e.g., Rippon 2000). Since then, these habitats
have been increasingly exploited, providing location for settlement, agriculture and harbours; source
of food, water and raw materials; and a focus for transport, trade and exploration (Rippon 1997,
361
Table 2 Estimates of actual cover and historical losses of coastal wetlands (and when possible salt marshes) for European countries (and eventual
additional regional information), main attributed causes of loss and known history of exploitation
Cover Cause Exploitation
Country (km2) Loss of loss history (yr) Regional data/additional information Additional references
Albania 150 2
>550–600 km since D
mid-1900s
Belgium 8.3 n.a. 2000 Limited settlement during Iron Age Rippon 2000
Bosnia- n.a. n.a.
Herzegovina
Bulgaria n.a. n.a.
Croatia n.a. n.a.

Cyprus n.a n.a.
Denmark 8851* 60% ^ since 1870 D Country’s coastline shortened by 1168 km (14%)
Estonia n.a. n.a
Finland 501 50% ^ D Losses of 22.8% ^ between 1950 and 1985
France 3 813 86% in twentieth D, U In Bretagne, 40% of coastal wetlands lost since 1960, and 66% of the EEA 2006a
362
century remaining seriously affected by drainage; ongoing losses in the
Languedoc Roussillon wetlands
Georgia n.a. n.a.
Germany 6809* 56.6% ^ in 1950s to D, U 2000 In the Wadden Sea, 200 km2 of salt marshes lost during 1950–1984, Dugan 1993, Rippon 2000,
1984 and only 400 km2 remain today; in the whole Wadden Sea (not only Reise 2005
Germany) 14,650 km2 of coastal wetlands lost since the eleventh
century (33% of salt marshes in 1930–1987)
Greece 1011 73% salt marshes in WQ, D EEA 2006a

twentieth century
Iceland n.a. n.a.
Ireland n.a. n.a. D >250 salt marshes are located/mapped; 65 km2 of estuarine area Curtis & Skeffington 1998,
reclaimed in the Shannon estuary and 20 km2 in the Wexford Harbour Healy & Hickey 2002
mainly in nineteenth century
Italy n.a. 25,000 km2 ^ since D, S, 2500 60% of the estimated losses occurred in 1938–1984; on the Po river Stanners & Bourdeau 1995,
<1000 of Roman times U, A delta, >70% of salt marshes reclaimed in 1870s–1960s; in Friuli Cencini 1998
salt 6000 km2 of salt Venezia Giulia 631 km2 lost in 1877–1990
marshes marshes in twentieth
century
Latvia 1426 n.a.
Lithuania 413 70% ^ during the past D
30 yr
Malta n.a. n.a.
Monaco n.a. n.a.
Montenegro n.a. n.a.
Netherlands 4043 (85 4000 km2 of salt D, WQ, >2500 Humans affected salt marshes since at least the Iron Age; earliest dam Wolff 1992, 1997, 2000,
of salt marshes between U dates 175 B.C. Rippon 2000
marshes) twelfth century and
second half twentieth
century
Norway n.a. n.a.
Poland n.a. 89.8% ^ D

Portugal 795 n.a. D, S, E 70% of salt marshes in the western Algarve lost before 1988 Allen 2003
Romania 5297 4000–6600 km2 ^
Russiaa 5786* n.a.
Slovenia n.a. n.a.
Spain 1041 60% D, U Most of the losses occurred in 1965–1990 EEA 2006a
363
Sweden 6000 23% ^ since early D >50% degraded since early 1800s; in some areas losses up to 80–90%
1800s
Turkey n.a. n.a.
Ukraine n.a. n.a.
United 5966 >50% of salt marshes D, U, E 2500 Coastal/marine wetlands comprise (in km2) 14 sand dune slack, 2658 Rippon 1997, 2000,
Kingdom (450 of since Roman times; 38 estuarine waters, 2793 intertidal flats, 450 salt marsh and 50 saline Davidson et al. 1991,
salt lagoons in the latter lagoons; salt marsh loss most significant in the 1800s but still ongoing Boorman 2003
marshes) half of 1980s; >913 (100 ha yr−1).
km2 of estuary area
Note: If not otherwise specified in the reference column, information is derived from Nivet & Frazier (2004) and references therein, while information about history of exploitation
is generally derived from Rippon (2000). A = aquaculture, D = drainage/embankment/land claim/conversion (e.g., to agriculture), E = erosion/sea-level rise/coastal squeeze, n.a. =
not available, S = altered sediment loads (e.g., from inland deforestation), U = urban/harbour developments, WQ = water quality degradation. Note that the definition of coastal
wetland is vague for most countries, and in many cases (indicated as *) only important or large marine wetlands were included. Concerning habitat loss, in many cases (indicated
as ^) estimates refer to total wetlands because no distinction was made between coastal and inland wetlands, and often no time span is indicated. Overall, estimates should be
considered as broad indications.
a Including Asian Russia.
Major complexes
Isolated sites >500 ha
Small <500 ha, frequent
Small <500 ha, scattered
Figure 7 Distribution of salt marshes along European coastlines. (From Boorman 2003. With permission.)
2000, Knottnerus 2005, Wolff 2005). European estuaries support major cities and harbours and
have an enormous economical and social importance. This long history of human exploitation has
deeply altered these habitats in extent and ecological characteristics. Although historical information
exists for some regions or localities (Table 2), knowledge of the extent of loss of coastal wetlands
is generally limited.
A first overview of E.U. wetlands and their loss was provided by Jones & Hughes (1993), and
more recently by Nivet & Frazier (2004), but they could not produce a European-wide estimate
because of the scarcity and poor quality of most data available. Denmark, the Netherlands, Germany,
Finland, Lithuania, the United Kingdom, Spain, Greece, Italy, France, Poland, Romania and parts
of Portugal and Sweden have reported losses of wetland exceeding 50% of original area, with peaks
above 80% for some regions (Table 2). These estimates sometimes refer to whole wetlands (without
distinction between coastal and inland) and most often cover a relatively small time span (over the
last century). However, when historical and archaeological documentation are available, it is evident
that significant losses of coastal wetlands started in Roman times and locally even earlier than that
(e.g., Wolff 1992, Allen 1997, Cencini 1998, Rippon 2000). Overall, it has been suggested that in
the Mediterranean Sea 28,000 km2 (>90%) of coastal wetlands have been lost since Roman times
(UNEP/MAP/PAP 2001). Recent estimates have also suggested that approximately two thirds of
all European coastal wetlands that existed at the beginning of the twentieth century have now been
lost (EEA 2006a).
364
Some of the best historical records of the loss of coastal wetlands are from the estuaries around
the United Kingdom, the Wadden Sea and the Po river delta in the north Adriatic Sea. In the United
Kingdom, coastal wetlands were already used in prehistoric times for salt production, pasture
and collection of wild resources (Rippon 2000). During Roman times a more systematic modifi-
cation of coastal marshes began, with documented reclamation of some areas around the Severn
estuary (Rippon 1997). Since mediaeval times, the use of coastal wetlands (e.g., for settlement,
agriculture and food) has increased continuously in intensity; salt marshes were systematically
enclosed, drained, settled and used for agriculture and pasture, leading to large-scale claims
throughout much of British estuaries (detailed historical reconstructions of these developments are
given by Rippon 2000). Overall, in the United Kingdom it is estimated that about 913 km2 of estuary
area and 550 km2 of salt marshes have been claimed since Roman times for agricultural, urban,
harbour and industrial developments (Davidson et al. 1991, Nivet & Frazier 2004). Decline was
most significant in the 1800s but it has continued until today. For example, in the Wash, 858 ha of
salt marshes were converted to agricultural use between 1970 and 1980 (U.K. Biodiversity Group
1999). Similarly, average losses of 20–25% of salt marshes, with individual sites suffering losses
of 10–44%, occurred between 1973 and 1998 in Kent and Essex (Cooper et al. 2001, Adam 2002),
where many salt marshes are ‘squeezed’ between an eroding seaward edge and fixed flood defence
walls (U.K. Biodiversity Group 1999, but see Wolters et al. 2005 for debate about the causes of
salt marsh erosion in southeast England). Overall, in England and Wales it is estimated that 15%
of salt marshes were lost between the 1940s and the 1970s (Nivet & Frazier 2004).
The history of human transformation of the Wadden Sea ecosystems has been recently reviewed
(Lotze et al. 2005). Similar to estuaries in the United Kingdom, exploitation of the wetlands in this
region started in prehistory, when people collected wild fauna and flora for subsistence, grazed
cattle and sheep and made the first attempts to transform the coastal landscape and control the flow
of water (Rippon 2000, Knottnerus 2005). Although these activities did not result in the disappear-
ance of salt marshes, they changed the composition of their vegetation considerably (Wolff 2000).
The conversion of coastal wetlands into arable land and freshwater lakes became systematic before
the eleventh century (Reise 2005), and after the introduction of windmill technology, in the sixteenth
century, it also became possible to drain shallow lakes (Wolff 2000). Embankments continued until
the second half of the twentieth century (e.g., Figure 4), supported by improved building technol-
ogies, and it was only after the 1960s that exploitation slowed and there was a growing focus on
conservation (Wolff 1997). Overall about 15,000 km2 of coastal wetlands have been lost during
this long history of progressive embankments (Reise 2005) and the whole Wadden Sea has been
reduced to nearly half of its primordial size.
Massive loss of coastal wetlands has also occurred in Italy. It is estimated that there were about
7000 km2 of coastal marshes remaining at the end of the nineteenth century, no more than 1920 km2
in 1972 and <1000 km2 today (Stanners & Bourdeau 1995). Compiled historical information is
limited to a few areas (e.g., the river Tevere; Keddy 2000). Cencini (1998) recently reconstructed
the evolution of the Po river delta in the north Adriatic Sea and the effects of human transformation
on coastal wetlands. Ancient sources and maps (e.g., from Pliny the Elder, Polybius and Strabo)
described the deltaic coastland as a continuous, almost impassable sequence of lagoon, marshes
and rivers. Since the Greco-Etruscan times, and more so after the consolidation of the Roman
Empire, the area was heavily inhabited and exploited, and important urban settlements, roads and
commercial ports were developed. Over the centuries, the main causes of transformation of the
delta were altered sedimentation patterns and direct land claim. Sediment loads were enhanced by
extensive inland deforestation carried out already by the Celts and Romans (Bondesan et al. 1995,
Barmawidjaja et al. 1995). Since the seventeenth and eighteenth centuries, hydraulic works of river
diversions, embankment and drainage took place, and after 1870 wetland drainage occurred sys-
tematically over large scales not only to improve hygiene conditions and eradicate malaria but also
365
Po River (main stream)

0 3 Km TA
EL
D
Po
Po di Goro
di G
GRANDE
1969
BONIFICAZIONE 1872
PO
noc
FERRARESE
RN
a c
1872 1930
DE
Po di Volano R.
MO
1878
Valle del Mezzano Valli Bertuzzi
1929
COMACCHIO
1958
Valli di Comacchio
1964
1931 Vene di Bellocchio
Valle
Santa
Po di
Primar
o R . - Reno R .
Punte Alberete
R.
Sillaro
1940 Pialasse ravennati
Reclaimed saltwater marshes RAVENNA

N
Existing saltwater marshes

C. Cencini, 1998
Reclaimed freshwater
R.
Existing freshwater marshes

Savio
Figure 8 Reclaimed (with indication of year of reclamation) and existing fresh- and saltwater marshlands in
the southern part of the Po river delta. (From Cencini 1998. With permission.) Overall 98% of the freshwater
marshes and >70% of the salt marshes were reclaimed between the 1870s and 1960s.
to claim new farmland. Reclamation ended by the 1960s with an almost-complete elimination of
marshes and irreversible changes to the hydrological and agricultural asset of the coastal plains.
Overall in the Po delta, 98% of the freshwater marshes and more than 70% of the salt marsh that
existed at the beginning of the twentieth century have been claimed (Figure 8). The ancient delta
(until the twelfth century A.D.) covered about 1300 km2, while the modern delta covers only 730 km2.
366
Trends and threats
Pollution, eutrophication, drainage, conversion to agriculture and aquaculture, changes in sedimen-

tation and hydraulic regimes, changes in sea levels, global warming, invasive species, fishery
overexploitation and disruptive fishing techniques, urban development, shipping, tourism and water
sports are generally considered the most serious threats to present-day coastal wetlands (e.g., Dugan
1993, Cencini 1998, Jansson & Dahlberg 1999, Wolff 2000, Kennish 2002, Hiscock et al. 2005).
However, limited information is available on the extent to which these activities affect and impair
specific coastal wetlands and thus the general importance in causing further habitat loss is difficult
to rate.
Although in some regions embankments and drainage have been stopped or even reversed (e.g.,
Wolff 1997, Cencini 1998), land loss still continues to be the most pervasive threat to coastal
wetlands and salt marshes. Between 1990 and 2000 there has been a total net loss of 390 km2 of
coastal wetlands around European coastlines, a significant proportion of which was lost as a result
of drainage to reclaim land for development and afforestation (EEA 2006a). Expanding aquaculture
and tourist infrastructures (e.g., marinas) are also making intensive use of estuaries and wetlands,
with limited consideration of the consequences on these systems (EEA 2006a).
Global changes in sea levels are beginning to pose severe threats to salt marshes, coastal
wetlands and river deltas because of the strong dependence of these habitats on water-level fluc-
tuations and tidal regimes (Adam 2002, Morris et al. 2002). It has been suggested that the projected
sea-level rise could cause the loss of up to half of European current coastal wetlands (EC 2004),
with some of the largest losses expected to occur around the Mediterranean and Baltic Seas (Nicholls
et al. 1999). When combined with other losses directly or indirectly due to human action up to
70% of the world’s remaining coastal wetlands could be lost within the next 100 yr (Nicholls et al.
1999), although there is considerable uncertainty. In the United Kingdom, for example, it has been
predicted that there could be a loss of freshwater and brackish habitats of around 4000 ha as a
consequence of the combined effects of sea-level rise and a temperature increase of 3–4°C (Lee
2001).
Such habitat losses from sea-level rise could naturally be compensated for by the habitat
regressing or developing inland. However, these inland areas are often developed and this coastal
infrastructure is defended by building barriers; marshes are thus caught in a ‘coastal squeeze’
between rising seas and expanding development (Doody 2004). In the United Kingdom, for exam-
ple, current ongoing losses of 100 ha yr−1 of coastal salt marsh have been estimated, due to
erosion/reduced sediment inputs/land subsidence and coastal defence measures to counteract ero-
sion (U.K. Biodiversity Group 1999, Hughes & Paramor 2004). Fortunately, since the 1990s a new
approach has been developed to address coastal erosion in Europe, and increasingly the sea is being
allowed to break through barriers and to re-create fringing habitats including salt marshes and
lagoons (e.g., EC 2004, Hughes & Paramor 2004, Kabat et al. 2005).
Protection measures
The importance of preserving coastal wetlands is being increasingly recognised in Europe (Jones &
Hughes 1993), where wetlands are now the target of numerous international and national initiatives
and regulations for conservation, wise use and restoration. Government commitments have been
encouraged though various initiatives, such as the Ramsar Convention and the Rio CBD (Table 1).
Atlantic, Mediterranean and Baltic salt marshes are specifically listed as habitats in Annex I of the
Habitats Directive, with some types of marshes identified as priority habitats for conservation, and
wetlands in general fall into several broad habitats of the directive, such as ‘Coastal lagoons’,
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‘Estuaries’ and ‘Large shallow inlets and bays’. Many coastal wetlands are also recognised as SPAs
according to the Birds Directive.
As a response to these initiatives, European states have developed their own national strategies
and action plans. As an example, in the United Kingdom approximately 80% of present salt marshes
(50% in northwest Scotland) are currently designated as Sites of Special Scientific Interest (SSSIs),
and in Northern Ireland five out of seven estuaries containing salt marshes have been designated
as Areas of Special Scientific Interest (ASSIs) (U.K. Biodiversity Group 1999). Ten areas in the
United Kingdom have been proposed as SACs (Habitats Directive), and 27 major salt marshes and
many smaller sites are included in SPAs (Birds Directive) and Ramsar sites: the target goal is to
stop further net loss of coastal wetlands, and restore 40 ha of salt marsh yr−1, to replace the 600 ha
lost between 1992 and 1998 (U.K. Biodiversity Group 1999).
Seagrass meadows
Seagrasses are rhizomatous, clonal, marine plants that form some of the most valuable and pro-
ductive coastal ecosystems in the biosphere (Costanza et al. 1997). They provide food and habitat
for a variety of biota and play a fundamental role in carbon and nutrient cycling, control of water
quality and sediment dynamics (Duarte 2002). Seagrasses can colonise a variety of coastal habitats
from estuarine to marine, subtidal to intertidal, sedimentary to rocky. Global area estimates for
seagrasses are beginning to be developed but at present there is no comprehensive dataset of actual
seagrass distribution, and even regional datasets can be limited. The World Atlas of Seagrasses
(Green & Short 2003) provides the most current and comprehensive compilation of information,
documenting some 177,000 km2 of seagrass and suggesting a tentative global acreage estimate of
500,000 km2.
Several seagrass species occur along the European coastline, including the natives Zostera
marina, Z. noltii, Ruppia maritima, R. cirrhosa, Cymodocea nodosa and Posidonia oceanica
(endemic to the Mediterranean Sea) plus Halophila stipulacea, which was recently introduced in
the Mediterranean Sea. Seagrasses around European coastlines are now increasingly well monitored
and published accounts exist for many sites and regions (Table 3). National inventories of seagrass
distribution are available (e.g., Davison & Hughes 1998 for the United Kingdom) or are being
prepared (e.g., REBENT programme for France). There is not, however, an organised comprehen-
sive inventory of the distribution and coverage of seagrasses in Europe, and records for some regions
are very incomplete. Furthermore, although in some regions or countries most seagrass beds are
known and located, their actual coverage has not been determined. The World Atlas of Seagrasses
(Green & Short 2003) documents a coverage of seagrasses of 1850 km2 in Scandinavia and the
Baltic Sea, 338 km2 in western Europe (United Kingdom, Wadden Sea, Portugal and Atlantic France
and Spain), 4152 km2 in western Mediterranean countries (Italy, France and Spain), and 950 km2
in the northwest Black Sea (Figure 9). It has been conjectured that seagrasses could be much more
abundant in the Mediterranean, covering from 25,000 to 45,000 km2 (Pasqualini et al. 1998).
Present-day seagrasses along European coasts are often described as in a degraded state (Green
& Short 2003), with low shoot densities, high mortality rates, and high fragmentation. This is the
case, for example, of seagrasses along the Mediterranean coasts of Spain (e.g., Table 3; Duarte
2002, Marbà et al. 1996, Delgado et al. 1997), including some deeper ones along the southeastern
coasts, where trawling damages up to 40% of the total Posidonia oceanica surface (Tudela 2004).
Along the Ligurian coast of Italy, P. oceanica meadows (50 beds covering 48 km2) have been
severely degraded due to coastal modification and town developments (Bianchi & Peirano 1995).
Some of these beds were severely damaged in the early 1990s by the wreck of the oil tanker Haven
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Table 3 Estimates of actual cover and historical losses of seagrasses for European countries (and eventual additional regional information)
and main attributed causes of loss
Country Cover (km2) Loss Cause of loss Regional data/additional information Additional references
Albania n.a. n.a
Belgium Not present
Bosnia- n.a. n.a.
Herzegovina
Bulgaria n.a. n.a.
Croatia n.a. n.a. I Considered as widespread; severe declines of Posidonia oceanica in Istria Zavodnik & Jaklin 1990,
between 1938–1998. Local invasions by Caulerpa taxifolia. Meinesz et al. 2001
Cyprus n.a. n.a. Considered as widespread.

Denmark 1380–1710 75–80% in WD, Sa, EU, Total cover is the sum of 30 km2 in the Wadden Sea and 1350–1680 km2 Reise 1994, Rask et al.
1900–1990s MG on the eastern coasts; vertical depth distribution reduced by 50% between 1999
1900 and 1996–1997 (from 5–6 to 2–3 m in estuaries and from 7–8 to
4–5 m in open waters); 261 km2 lost in Limfjiorden between 1900 and
1994 and 559 km2 lost in Oresund between 1900 and 1996–2000; local
369
losses continue nowadays (e.g., in 1994 eelgrass temporarily lost at Funen
Island due to summer anoxia).
Estonia n.a. n.a.
Finland <10 No losses since Seagrasses presumably not affected by the wasting disease.
1968
France n.a. n.a. WD, LC, P, I, Along the Atlantic coasts >70 sites mapped and coverage of many known Meinesz et al. 1991,
1150 km2 of 30–40% of F, T (most beds are 1–5 ha, but at least 10 beds cover 10 to >100 ha). South 2001, Glemarèc et al.
P. oceanica P. oceanica in of Arcachon there is the largest Zostera nolti bed in Europe (70 km2 in 1997, Duarte 2002,
recent decades 1984) and a large bed of Z. marina (4 km2 in 1984). The Glenan EEA 2002
Archipelago lost 6 km2 of seagrasses during 1932–1992. In the
Mediterranean many local documented losses of P. oceanica (e.g., close
to Marseille, 5–6% per decade between 1900 and 1970, nowadays 90%
is deteriorated; along the French Riviera, 30.57 km2 lost since 1800;
seagrasses virtually extinct from the region of Toulon). 3184 ha affected

by the invasion of C. taxifolia.
Table 3 (continued) Estimates of actual cover and historical losses of seagrasses for European countries (and eventual additional regional
information) and main attributed causes of loss
Country Cover (km2) Loss Cause of loss Regional data/additional information Additional references
Georgia n.a. n.a.
Germany 170 n.a. WD, EU, S, Most seagrasses occur in Wadden Sea; only present on eastern coasts. Baltic Reise 1994, Bartsch &
MGb, LC Sea: in Kiel Bight decreased from 6 to 2 m in depth between 1960 and Tittley 2004
end of 1980. No losses in Greisvald Lagoon between 1930 and 1988.
North Sea: Z. marina extinct at Helgoland island as early as 1928.
Greece n.a. n.a. Considered as widespread.
Iceland n.a. n.a.

Ireland n.a. n.a.
Italy n.a. 30–40% of S, EU, P, I, T Most beds are mapped, and coverage of many is known (estimated Barmawidjaja et al.
P. oceanica in 2350 km2 in Liguria, Lazio, Sardinia, Veneto and Friuli). Estimate of loss 1995, Piazzi et al.
recent decades is derived from estimates from France. P. oceanica virtually extinct in the 2000a,b, Guidetti 2001,
north Adriatic Sea, in some areas since earlier than 1850s. Z. marina Meinesz et al. 2001,
370
disappeared from areas of the Venice lagoon. 9414 ha affected by the Milazzo et al. 2004
invasion of C. taxifolia.
Latvia n.a. n.a.
Lithuania Extinct? 100%? MGc Considered abundant over thousands of hectares in the past.
Malta n.a. n.a.
Monaco n.a. n.a. I Severe invasion by C. taxifolia. Meinesz et al. 2001
Montenegro n.a. n.a.
Netherlands 200 n.a. WD, Ec, S, Most seagrasses are in southwest estuaries; in the Grevelingen estuary, the Reise 1994, Short &
Virtually extinct EU, Fd construction of two dams facilitated the growth of Z marina (from 12 km2 Wyllie-Echeverria
in Wadden Sea in 1964 to 34 km2 in 1985) and the disappearance of Z. noltii, which 1996, Wolff 2000, 2005
before was the most common seagrass (large-scale, unexplained die-off
of Z. marina since 1986–1987). In the Wadden Sea 145 km2 lost in
1919–1971 and 3 km2 since) and only 1–2 km2 are left nowadays.
Norway n.a. No losses since
1930
Poland n.a. n.a. MGc, EU Virtually extinct in Puck Lagoon in 1957–1987, with subsequent
recolonisation in some areas.
Portugal n.a. n.a.
Romania n.a. n.a.
Russia n.a. n.a.
Slovenia n.a. n.a.
Spain n.a. 30–40% of S, F, I, A, T Estimate of loss is derived from estimates from France. In the Marbà et al 1996,
>1000 in P. oceanica in Mediterranean, 57% of P. oceanica beds were under regression in 1996 Meinesz et al. 2001,
Mediterranean recent decades along 1000 km of coasts, and some are now extinct, with losses Duarte 2002, EEA 2002
concentrated over 400 km of coasts. 58% of P. oceanica beds and 52%
of seagrasses are degraded along the Catalan coasts and near Alicante,
respectively. Local invasions by C. taxifolia.
Sweden n.a. n.a. EU, MG At least 60–130 km2 along the southern Baltic coasts. Losses of 58% Baden et al. 2003
(10.61 km2) of seagrasses in the Skagerrak in 1980s–2000.
Turkey n.a. n.a. Considered as widespread.
Ukraine n.a. n.a. 950 km2 in the northwestern bays.

United n.a. n.a. WD Most beds mapped, and coverage of many known. About 140 sites of Davison & Hughes 1998,
Kingdom Z. marina and about 70 sites of Z. noltii, covering from 12 km2 (Cromarty U.K. Biodiversity
Firth) to 20–40 hae. The Maplin Sands hosts one of the largest surviving Group 1999
population of Z. noltii in Europe (325 ha). Before 1900s, seagrasses were
common; they severely declined in 1920s–1930s and have not recovered
371
yet, particularly in southern and eastern England.
Note: If not otherwise specified in the reference column, information is derived from the World Atlas of Seagrasses (Green & Short 2003) and references therein. A = aquaculture,
E = engineering works and embankments, EU = eutrophication, F = fisheries, I = invasive species, MG = growth of ephemeral macroalgae (often a consequence of eutrophication),
LC = land claim/waterfront development, n.a. = no comprehensive estimate available, P = urban and/or industrial pollution, S = increased water turbidity/load of sediment, beach
replenishments, T = tourism, WD = wasting disease. Note that the estimates of covers should be regarded as minimal representation of the actual coverage in most cases. Also
note that most often time span is not indicated.
a Decrease of maximum Secchi depth from 12 m in 1900 to 6 m in the 1990s.
b Seagrasses replaced by filamentous algae. This may not necessarily be the cause of seagrass loss.
c Including the closure of the Zuidersee in 1932.
d Eelgrass harvesting until 1930, shell fisheries after 1970.
e Other important sites are the Exe Estuary, the Solents marshes and the Isles of Scilly, Morfa Nefyn, Milford Haven, the Moray Firth, Carlingford Lough, Dundrum Bay,
Strangford Lough and Lough Foyle.

0 250 500 1,000

Km
Figure 9 Map of the distribution of seagrasses in Europe. (Data courtesy of UNEP World Conservation
Monitoring Centre.) The map should be considered indicative of general distribution not areal extent.
(Sandulli et al. 1994). The extent of seagrass degradation is, however, difficult to quantify even
locally.
There is increasing awareness about the severe degradation of seagrass meadows (e.g., see, among
others, the reviews by Short & Wyllie-Echeverria 1996, Hemminga & Duarte 2000, Duarte 2002).
Reports consistently identify a long-term trend of worldwide seagrass decline, about 70% of which
can be probably attributed to direct human-induced disturbance (Short & Wyllie-Echeverria 1996).
Less information is available concerning the degradation caused by indirect impacts (Duarte 2002).
It has been estimated that a global loss of 12,000 km2 occurred during the 1990s alone (Short &
Wyllie-Echeverria 1996), which represents about 7% of the known distribution of seagrasses (Green
& Short 2003). Data covering longer time spans are rare. Based on data from 12 temperate estuaries
around the world, it has been estimated that over time these systems may have lost approximately
65% of their seagrasses (Lotze et al. 2006). No comprehensive organised historical information
seems to be available for Europe and information is limited to restricted areas (Table 3).
There are different trends for seagrass losses in northern and Mediterranean European countries
(Green & Short 2003). In northern Europe, before the early 1900s, several seagrass species,
including Zostera marina, were common. They were harvested for a variety of uses, including use
for fuel, packing, upholstery, insulation, roof material, filling of mattresses and cushions, feeding
and bedding for domestic livestock, fertiliser and as resource to obtain salt. Their abundance was,
however, severely reduced in the 1930s by a ‘wasting disease’, caused by the pathogenic slime
mould Labyrinthula zosterae (e.g., Den Hartog 1987). The disease led to the catastrophic die-back
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of eelgrass (Zostera marina) meadows along the north Atlantic coasts, with loss of almost 90% of
the Zostera populations in north Atlantic western Europe. Some beds progressively recovered, but
substantial areas remain lost from most beds.
There is uncertainty about the causes of this disease outbreak and there has been much debate
about whether concomitant human impacts that already weakened the plants contributed to the
outbreak (Den Hartog 1987). The decline particularly affected sublitoral beds, while intertidal
populations were less affected. Probably the best records of the disruptive effects of the wasting
disease are from Denmark, where records of eelgrass distribution date back to 1900 (Boström et al.
2003). In 1900 eelgrass covered about 6726 km2 (Figure 10); by 1940, 93% of the distribution of
vegetated areas was lost. Since 1960, there has been a slow recolonisation. Currently, there is
approximately 20–25% of the distribution recorded in 1900 (i.e., a total loss of 75–80%). The
greatest loss has been in deep Zostera populations and in Denmark the vertical depth distribution
of Z. marina was reduced by about 50% during the twentieth century, from a historical depth limit
of 5.6–11 m to a recent limit of 2.5–8 m in sheltered and exposed areas, respectively (Hemminga &
Duarte 2000, Baden et al. 2003).
Similar dramatic losses are described for the United Kingdom (Davison & Hughes 1998 and
references therein) and the Wadden Sea (Reise 1994, Wolff 2000, Lotze 2005). The distribution of
seagrasses in the United Kingdom was only systematically described in the 1930s after the outbreak
of wasting disease, when Z. marina was already scarce and restricted to few sheltered lagoons but
there are indications that seagrasses were widespread until 1917 (Davison & Hughes 1998). There
is some uncertainty about when recovery started. According to some, recovery began in 1933 and
was quite rapid, with some beds fully recovering within a few years of the 1930s epidemic, while
according to others the disease continued to affect Zostera populations until the mid-1940s and
recovery did not really begin until the 1950s. Nowadays, most Zostera beds have not fully recovered,
particularly in southern and eastern England where the species was once abundant, and only 20 of
Britain’s 155 estuaries have eelgrass meadows >1 ha in extent (Davison & Hughes 1998).
Before the 1930s the Wadden Sea also contained large subtidal, seagrass beds. These have been
exploited since historical times for construction and insulating material and to fill mattresses and
cushions. In the Dutch Wadden Sea, from the thirteenth century to 1825, eelgrass was used to build
dykes. The construction of 1 m of seawall required about 8–20 m3 of compacted eelgrass, equivalent
to about 40–100 m3 of fresh eelgrass, which in some years corresponded to about 1–10% of the
annual production (Wolff 2005). Decline of seagrasses appears to have occurred over two phases
(Reise 1994): one acute in the 1930s, caused by the wasting disease, after which most subtidal
eelgrass beds did not recover, and one more gradual that began in the 1960s, mostly driven by
eutrophication. These declines first affected subtidal eelgrass beds, and subsequently also intertidal
ones, leading to the almost extinction of seagrasses in some regions of the Wadden Sea (e.g., the
Dutch Wadden Sea, where cover dropped from 150 to 1–2 km2; see Table 3), and to the disappear-
ance of numerous species associated with seagrasses (Wolff 2000).
Along Mediterranean coasts, reliable estimates made by direct observation of the area of
seagrass lost or degraded are limited (Green & Short 2003). It is estimated that in the past Posidonia
oceanica meadows may have covered 50,000 km2 in the whole basin (Duarte 2002), which con-
sidering present estimated covers of seagrasses in the Mediterranean and Euro-Asian Seas (Green
& Short 2003) would make an overall loss >85% (but probably many existing seagrass meadows
are not presently documented). Rapid local regression (up to complete disappearance) of P. oceanica
meadows is known to have occurred at numerous localities in France, Italy and Spain (Table 3). It
is estimated that shoot density of P. oceanica in the western Mediterranean has decreased by up
to 50% over a few decades, with major losses between 10 and 20 m depth (EUCC 1998). For the
French mainland coast, overall habitat loss is estimated as about 10–15%, which would increase
up to 30–40% if the decline in shoot density is also taken into account. Overall, these figures are
373
North Sea 1901
Sweden
Kattegat
Denmark
∅resund
Baltic Sea
Germany
North Sea 1941
Sweden
Kattegat
Denmark
∅resund
Baltic Sea
Germany
North Sea 1994
Sweden
Kattegat
Denmark
∅resund
Baltic Sea
Germany
Figure 10 Area distribution of eelgrass Zostera marina (in dark grey) along Danish coasts in 1901, 1941 and
1994. (Modified after Boström et al. 2003. With permission.) In the 1930s, eelgrass populations were severely
affected by a wasting disease, and in 1941 they covered only 7% of the areas occupied in 1901. Recolonisation
took place after the 1960s, but in 1994 cover was still only 20–25% of that in 1901.
374
considered a good estimate for most western Mediterranean coastlines, with notable exceptions
around the islands and in the eastern Mediterranean (EUCC 1998).
There have been clear losses of seagrasses on the Italian coast of the north Adriatic Sea.
Geological data have shown that seagrass beds were probably common before the 1800s and
experienced dramatic regressions to virtual extinction in the last two centuries (Barmawidjaja et al.
1995, Caressa et al. 1995, Rismondo et al. 1997). For example, faunal changes and the sudden
disappearance of epiphytic foraminiferans in a sediment core in front of the Po river delta suggest
that seagrass beds were present up to 1840 in this area, and that increased load of fine sediment
and nutrients between 1840 and 1870, due to substantial changes to the main outflow canals of the
Po river, was probably the main cause of their sudden disappearance (Barmawidjaja et al. 1995).
Similarly, dead beds of P. oceanica have been found at several sites about 8 miles offshore from
the Venice lagoon, indicating the likely past presence of P. oceanica (Rismondo et al. 1997). In the
Gulf of Trieste the regression has been more recent and there has been some direct documentation.
P. oceanica was reported as common in the Gulf of Trieste at the beginning of the 1900s (Caressa
et al. 1995). In 1938 the species had declined but was still present all over the coastlines of the
Istrian peninsula, while just about 30 yr later only the Koper meadow was detected. Nowadays,
P. oceanica is present only in a fragmented meadow along the coastline of Koper (Slovenia, at the
southern side of the Gulf of Trieste) and in a very small area along the coasts of Grado (on the
Italian side of the Gulf of Trieste). This drastic reduction has been attributed to a steep increase in
water pollution during the last 50 yr as a consequence of industrial and harbour development in the
Gulf of Trieste (Caressa et al. 1995).
Another case of rapid regression of P. oceanica meadows is in the French Riviera, from Menton
to the Rhône delta (Meinesz et al. 1991). Intensive waterfront development started around 1800
covering natural coastal habitats with recreational harbours, artificial beaches, landfills (for the
tourist industry) and large commercial and military complexes, ports and airports. A total of 185
reclamation projects ‘occupied’ 106 km (16.2%) of the coastline, directly removing 30.57 km2 of
bottom substrata and greatly affecting surrounding areas (by modifying water movements and sedi-
mentation patterns). Overall a total of 9.7% of the shallow water zone between 0 and −20 m and
14.5% of the zone between 0 and −10 m were irreversibly destroyed. The vast majority of this area
was occupied by P. oceanica meadows, which were estimated to originally cover a total of 200 km2.
Trends and threats
Many anthropogenic factors are considered responsible for the ongoing degradation and decline of
seagrasses in Europe as well as globally (e.g., see, among others, the reviews by Short & Wyllie-
Echeverria 1996, Davison & Hughes 1998, Hemminga & Duarte 2000, Duarte 2002, Green &
Short 2003). The most important of these threats is likely to be poor water quality from pollution,
eutrophication and excess sedimentation. These impacts are associated with, and enhanced by,
urban and tourist waterfront developments, port constructions, beach replenishments and other
interventions for shoreline stabilisation.
Severe seagrass loss is still in progress in Europe, as evident along 200 km of the Skagerrak
Swedish coast (Baden et al. 2003). Here, 50 of 69 mapped meadows of Zostera marina have shown
average declines of 58% between the 1980s and 2000, corresponding to a lost surface of about
1061 ha. Most declines are related to a reduction of the upper and the lower depth distribution of
seagrasses, resulting in narrower meadows, but in some areas seagrass meadows have disappeared
completely, with dramatic effects on the fish assemblages (Pihl et al. 2006). The reasons for this
continuing loss of seagrasses are not known but could be related to an excess growth of phytoplank-
ton and filamentous or other ephemeral macroalgae as a consequence of eutrophication. These
micro- and macroalgae outcompete seagrasses (Hauxwell et al. 2001) and decompose on the bottom,
375
favouring anoxia events like those that caused the recent temporary disappearance of Z. marina in
1994 around the island of Funen in Denmark (Rask et al. 1999).
There is also current seagrass loss in the Mediterranean Sea as a consequence of the invasion
of Caulerpa taxifolia (Meinesz et al. 2001, but see Jaubert et al. 2003). This species competes for
space and resources with the seagrass Cymodocea nodosa (Ceccherelli & Cinelli 1997) and is
thought to be able to damage Posidonia oceanica beds, particularly when these are already under
stress (e.g., de Villèle & Verlaque 1995).
Protection measures
Green & Short (2003) report that worldwide there are only some 247 MPAs that are known to
include seagrasses, spread in 72 countries and territories. This estimate may be conservative but
these authors note that this is far smaller than the number of MPAs with coral reefs (more than
660) or mangrove forests (over 1800). Even for these 247 MPAs, there are questions about their
effectiveness in protecting seagrass ecosystems particularly from threats such as poor water quality
(e.g., Marbà et al. 2002, Milazzo et al. 2004).
In Europe, numerous initiatives arising from the Rio CBD, the Habitats Directive and the Birds
Directive (Table 1) have led to seagrass meadows being specifically targeted for conservation and
restoration. Seagrasses are a named component of several habitats in the Habitats Directive,
including ‘Coastal lagoons’ (a priority habitat), ‘Sandbanks slightly covered by seawater all of the
time’, ‘Large shallow inlets and bays’, ‘Estuaries’ and ‘Mudflats and sandflats not covered by
seawater at low tide’ (EC 2003). Furthermore, Posidonia oceanica beds are a priority habitat in
Annex I of the Habitats Directive. International concern about the conservation of seagrass beds
has also led to the banning of trawling on seagrasses in EC waters (Tudela 2004).
As a response to these initiatives, European States have also developed national strategies and
initiatives. The U.K. Biodiversity Action Plan, for example, includes a Habitat Action Plan for
seagrass beds (U.K. Biodiversity Group 1999). Accordingly, areas of seagrass are included in some
coastal ASSIs/SSSIs, Ramsar sites, SPAs and voluntary MPAs. Two of the three U.K. Marine Nature
Reserves have seagrass beds and the habitat occurs in a number of areas proposed as SACs.
Macroalgal beds
For the purpose of this review ‘macroalgal beds’ refer to kelps, fucoids and other complex, erect
brown and red macroalgae that produce relatively large biogenic habitats. Macroalgal beds form
diverse, productive and valuable temperate coastal ecosystems (Steneck et al. 2002). They are wide-
spread on shallow hard substrata around Europe (Birkett et al. 1998b, Steneck et al. 2002, Thibaut
et al. 2005 and references therein), including rock, boulders, cobbles and human-made structures
from the intertidal down to more than 30 m in depth. Laminaria and Fucus are the main genera
along the coasts of northwest Europe, while Cystoseira and Sargassum are the main genera in the
Mediterranean Sea.
The macroalgal flora of the European coasts are among the best known and studied. Further-
more, large macroalgae have been the target of ecological studies for over a century and there is
extensive literature on physical and biological factors operating in these habitats (see, among many,
Lüning 1990, Ballesteros 1992). Surprisingly, however, quantitative information on the distribution
of macroalgal beds is limited and their extent is unknown. At present, there do not appear to be
comprehensive inventories for the macroalgae of any European country and it is difficult to estimate
the areal coverage even at regional or local scales. Distribution maps of rocky coast biotopes,
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including kelps, are available for the United Kingdom (Connor et al. 2004), and ongoing inventories
by the Joint Nature Conservation Committee (JNCC) are likely to result in estimates of the cover
of macroalgal beds for this country, at least for intertidal habitats, while little information is available
for the subtidal. Intensive research on kelp distribution, ecology and effects of human harvesting
has also been carried out along Norwegian coasts, where it is estimated that the dominant kelp
Laminaria hyperborea might cover between 5,000 and 10,000 km2 (Jensen 1998). Limited quan-
titative information on the distribution of macroalgal beds is available at a few regional or local
scales (e.g., the Albères coast, Thibaut et al. 2005; Kiel Bay, Vogt & Schramm 1991; the Öregrund
archipelago, Eriksson et al. 1998; the Gullmar Fjord, Johansson et al. 1998).
A perceived worldwide decline of macroalgal beds has been reported during the last decades
(Steneck et al. 2002, Airoldi 2003). Such decrease most often appears to be parallelled by a trend
of increasing abundance of turf-forming, filamentous or other ephemeral algae (e.g., Munda 1993,
Eriksson et al. 1998, Benedetti-Cecchi et al. 2001, Lotze 2005) that, once established, often inhibit
recolonisation of canopy-forming algae and other organisms (Airoldi 1998). Canopy-forming algae
and turfs have been suggested to represent alternative states in shallow temperate rocky reefs under
different disturbance and stress regimes (Worm et al. 1999, Airoldi 2003, Connell 2005). Macroalgal
beds have also been replaced by coralline dominated ‘urchin barrens’, where outbreaks of urchins
may have been the primary cause of macraolgal extirpation (Hagen 1995, Steneck et al. 2002,
Guidetti et al. 2003), or by mussel beds (Thibaut et al. 2005).
In Europe, almost no information on trends in abundance of macroalgae is available before the
1900s. In the twentieth century, conspicuous losses, sometimes to virtual local disappearance, of
complex macroalgae have been documented for coastal areas in several countries, including Iceland,
Norway, Britain and Ireland (Hagen 1995, Steneck et al. 2002 and references therein); Sweden
(Lundälv et al. 1986, Eriksson et al. 1998, 2002, Johansson et al. 1998, Nilsson et al. 2004);
Denmark (Middelboe & Sand-Jensen 2000); Finland and Germany (Kangas et al. 1982, Messner
& von Oertzen 1991, Vogt & Schramm 1991, Schories et al. 1997); Lithuania (Olenin & Klovaité
1998); Italy (Sfriso 1987, Cormaci & Furnari 1999, Benedetti-Cecchi et al. 2001, Guidetti et al.
2003, L. Airoldi unpublished data); France and Spain (Rodríguez-Prieto & Polo 1996, Thibaut
et al. 2005 and references therein); Croatia (Munda 1993, 2000) and Romania (Zaitsev 2006). The
causes of these losses are various but mainly include outbreaks of sea urchins and decreased water
quality as a consequence of pollution and/or enhanced sediment loads. These trends have been
generally traced through comparisons with historic floristic records, which are difficult to translate
into an estimate of the extent of habitat loss. Furthermore, complex native species were often
replaced by simpler macroalgae or non-native species, possibly affecting the status and functioning
of the systems but not their extent.
Frequently, the declines of macroalgae have been greater at depth so that their depth range has
become shallower (Lumb 1990, Messner & von Oertzen 1991, Bokn et al. 1992, Munda 1993,
Eriksson et al. 1998, 2002, Pedersén & Snoeijs 2001, Thibaut et al. 2005). In Kiel Bay (Germany,
western Baltic), for example, there has been a >90% decline in the biomass of Fucus spp. between
1950 and 1988, from about 40,000–45,000 t wet wt down to only 2400 t wet wt (Vogt & Schramm
1991). In the 1950s, Fucus spp. were the dominant macrophytes down to 6 m in depth and were
still frequent in the 1970s, whereas at the end of the 1980s, the species were not found at depths
>2 m. In the Öregrund archipelago (Sweden) the average depth penetration of F. vesiculosus
decreased significantly by 2 m between 1943 and 1996 and this species had completely disappeared
at depths >8 m (Figure 11). The amount of macroalgal habitat lost as a consequence of this depth
reduction has not been estimated (B.K. Eriksson personal communication) but it must have been
377
Max. depth Mean depth

0
∗
2
Depth (m)
6
8
1943–44
10 1984
1996
12
Figure 11 Changes in the depth (maximum and mean) distribution of Fucus vesiculosus in 1943–1944, 1984
and 1996 in the Öregrund archipelago. (From Eriksson 2002, based on Kautsky et al. 1986, Eriksson et al.
1998 and Eriksson & Bergstrom 2005; courtesy B.K. Eriksson.) *No mean depth data were available for 1984.
Data are averages ± 1 standard error. Maximum and mean depth penetration of F. vesiculosus decreased
significantly both between 1943–1944 and 1984 and between 1943–1944 and 1996 (t-test, n = 5 sites, p < .05).
considerable because quantitative measures of the distribution of F. vesiculosus indicated that this
species covered on average 20–50% of the substrata. Similarly, at Stora Bornö Island, on the
Swedish Skaggerak coast, the depth distribution of macroalgae declined on average by 2.8 m
between 1941 and 1998 (Eriksson et al. 2002). This loss was particularly severe for large, complex
macroalgae (>50 cm), which showed up to 8-m reductions in their depth distribution (Figure 12).
These complex macroalgae were replaced by simpler, thin filamentous and sheet-like forms.
Losses to virtual extinction of macroalgal species have been reported from other regions in the
Wadden Sea, southern France, the Venice lagoon and the Black Sea, for example. In the Wadden
Sea, at least 10 species of macroalgae have become extinct during the past 2000 yr, probably
because of the transformation of brackish waters into fresh waters and the destruction of native
eelgrass beds and oyster reefs that provided solid surfaces for attachment (Wolff 2000). Along the
1941 1998
2
4
6
Depth (m)
8
10
12
14
16
300 200 100 0 100 200 300
Estimate of cover (%)
Figure 12 Depth distributions of macroalgae with different thallus shape (thin filamentous and sheet-like =
white bars; coarse filamentous and thick, leathery algae = dotted bars) in 1941 and 1998 at Stora Bornö Island.
(Based on data from Eriksson et al. 2002. Courtesy B.K. Eriksson.) Bars are mean percentage cover in each
depth interval pooled for two vertical profiles (error lines are not shown for clarity).
378
Albéres coasts (southern France) dramatic reductions in abundance to extinction of populations of

Fucales (Cystoseira spp. and Sargassum spp.) occurred between the early 1900s and 2003 (Thibaut
et al. 2005). Only 9 of 14 species of Fucales documented in 1912 were present in 1978, with the
genus Sargassum entirely lost, and only 5 species were found in 2003. Seven of the extinct species
were considered frequent to abundant in 1937. Of the 5 species remaining in 2003, only 1 did not
show signs of regression. In the Venice lagoon, some 141 algal species were documented in 1938,
116 in 1962, 107 in 1987 and 96 in 1991 (Sfriso 1987, Sfriso & La Rocca 2005). Some of the
most damaged species in the lagoon were previously dominant large brown algae, including species
of Cystoseira, Fucus virsoides, and Sargassum hornschuchii. These algae were adversely affected
when channel excavations limited water exchange, leading to an increase of nutrient levels and
eutrophication within the lagoon, and in the 1980s their abundance dramatically decreased, some-
times to complete and permanent disappearance. These complex macroalgae were replaced by
ephemeral species, mainly green algae, and invasive species. In the last 10 yr, water quality has
improved in the Venice lagoon, and recent sampling detected an increase in the number of macro-
algae (Sfriso & La Rocca 2005), in part also linked to the introduction of several invasive species,
including the now-abundant large brown algae Undaria pinnatifida and Sargassum muticum. Along
the coasts of the Black Sea, several complex macroalgae (including brown algae of the genera
Cystoseira and Phyllophora) have virtually disappeared during the past 30 yr (Zaitsev 2006).
Estimates at some Romanian localities indicate that in 1971 these species attained considerable
biomasses (about >10,000 t fresh wt).
Trends and threats
Several factors are thought to be responsible for the continuing decline of kelps and canopy-forming
macroalgae and to pose serious threats to the future of rocky reefs in general (Steneck et al. 2002,
Thompson et al. 2002). Urbanisation is thought to have the most disrupting effects on kelps and
other canopy-forming algae, particularly by affecting water clarity and quality as well as other
habitat-related changes (e.g., Vogt & Schramm 1991, Munda 1993, Eriksson et al. 1998, 2002,
Benedetti-Cecchi et al. 2001).
In some northern European regions, including the west coast of Norway, the French channel
coast and parts of the U.K. coast, harvesting is also an issue (e.g., Christie et al. 1998). Kelps
washed up on the shore have been traditionally collected for centuries in some regions, for use as
an agricultural fertiliser and to improve the soil structure. Nowadays kelps and fucoids are harvested
from living beds to be used as basic resource in the alginate industry to produce emulsifying and
gelling agents. The most commonly harvested species include Laminaria hyperborea, L. digitata,
Ascophyllum nodosum and Fucus spp. with 70,000–80,000 t of seaweeds collected each year around
the coasts of both Brittany (Birkett et al. 1998b) and Norway (EEA 2002). Modern methods of
kelp harvesting (e.g., by trawling) seem to have a significant direct influence on kelp biotopes
(Birkett et al. 1998b, Christie et al. 1998).
Reef habitats and associated macroalgal beds are also severely damaged by disruptive fishing
techniques. For example, the collection of the date mussel Lithophaga lithophaga by use of hammers
and chisels, pneumatic hammers and explosives is still a widespread practice in most Mediterranean
countries, despite its legal ban. This practice directly and irreversibly destroys the rocky environ-
ment, causing the loss of canopy-forming seaweeds and the formation of barrens (Fanelli et al.
1994, Guidetti et al. 2003).
The introduction of harbour piers, jetties, dykes, seawalls, coastal defences and other armoured
artificial structures has in some regions led to an expansion in the distribution of native and non-
native macroalgae and other rocky-bottom species (Moschella et al. 2005). In the Wadden Sea, for
example, where hard substrata were naturally scarce, about 730 km of artificial structures have
379
introduced about 2–4 km2 of hard substrata, providing new habitats for a variety of rocky-bottom
species, including Laminaria saccharina (Reise 2005). Along the north Adriatic shores, which are
naturally devoid of rocky substrata, >190 km of rock-armoured structures (Figure 5), built mainly
in the past 40 yr (Bondesan et al. 1995), have introduced about 1 km2 of artificial hard substrata
within natural sandy depositional environments, which are now extensively colonised by the non-
indigenous, invasive canopy-forming macroalga Codium fragile ssp. tomentosoides (Bulleri &
Airoldi 2005). The extent of hard coastal structures is expected to increase in the future, with
profound but overlooked ecological consequences on native coastal environments (Airoldi et al.
2005).
Protection measures
Although kelp beds and other macroalgal habitats are not specifically targeted in the Habitats
Directive, species of the genus Fucus, Laminaria and Cystoseira and other macroalgae are named
components of ‘Reefs’ habitat (EC 2003). Other European initiatives also include the protection
of some species of complex macroalgae. For example, six Mediterranean species of the genus
Cystoseira and two species of Laminaria are listed in Annex I of the Bern Convention. The Action
Plan for the Conservation of Marine Vegetation in the Mediterranean Sea, adopted within the
framework of the Barcelona Convention, identifies the conservation of Cystoseira belts as a priority.
Several complex brown macroalgae are listed in the Red Books of Mediterranean and Black Seas
as endangered (e.g., Boudouresque et al. 1990, Zaitsev 2006). Furthermore, Lithophaga lithophaga
is included in Annex IV of the Habitats Directive and its collection is banned in most Mediterranean
countries to protect rocky reefs and associated macroalgal beds from the destructive consequences
of the fishery for this rather abundant date mussel (Russo & Cicogna 1991).
There are also national initiatives. For example, the commercial harvesting of kelps is strictly
regulated in Norway and in Brittany (Birkett et al. 1998b), including a system of rotation of
harvested areas introduced by the Norwegian government to ensure that each area of kelp forest
is harvested only once every 4 yr.
Biogenic reefs: oyster reefs

The native European flat oyster (Ostrea edulis) is a sessile, filter-feeding, bivalve mollusc that used
to be very abundant throughout its range (Korringa 1952). It is associated with highly productive
estuarine and shallow coastal water habitats with sediments ranging from mud to gravel. The natural
distribution of O. edulis is along the European Atlantic coasts from Norway to Morocco and across
the coasts of the Mediterranean and Black Seas. Their abundance declined significantly during the
nineteenth and twentieth centuries and wild native beds were considered scarce in Europe as early
as the 1950s (Korringa 1952, Yonge 1966, Mackenzie et al. 1997).
Remains of wild native oyster beds still occur in various regions, including the rivers and flats
bordering the Thames Estuary, the Solent, River Fal, the west coasts of Scotland and Ireland
(Kennedy & Roberts 1999, U.K. Biodiversity Group 1999, Tyler-Walters 2001, Jackson 2003), the
western part of the Swedish Kattegat region of the Baltic (Lozan 1996), the Limfjord region of
Denmark (Korringa 1952), the Adriatic Sea, where O. edulis is still captured in the wild (Barnabe
& Doumenge 2001), the Mar Menor (Spain), where a large flat oyster population, estimated at over
100 million individuals, still produces large amounts of spat (Cano & Rocamora 1996), and areas
of the Black Sea, where the species was still valuable commercially until the 1970s (Zaitsev 2006).
Limited information, however, is available about the current status of these oyster reefs and there
380
is debate about whether the fragmented patches of wild oyster habitats are self-sustaining or owe
their survival to the inputs of larvae from cultivated oysters (Korringa 1952).
Nowadays, aquaculture provides the main supply of native oysters in most European countries
(Mackenzie et al. 1997, Ocean Studies Board 2004). This industry has also been seriously affected
by epidemic diseases in recent decades, with documented losses of commercial stocks above 80%
in France (Kennedy & Roberts 1999, Ocean Studies Board 2004), and most Mediterranean native
oyster beds are in such poor conditions that they are unable to support intensive culture (Barnabe
& Doumenge 2001). Although marketplace demand for native oysters remains strong, the introduced
Pacific oyster Crassostrea gigas, which is easier to cultivate than the native oyster, now provides
the major share of oyster production in Europe (Cano & Rocamora 1996, Kennedy & Roberts
1999, Lotze 2005; see Figure 13C).
There is some documentation on the declines and loss of native oyster reefs, mostly from fishery
landing records; direct quantitative data are uncommon. In Europe oysters have been an extremely
popular food for centuries (Jackson 2003). Both ancient Greeks and Romans highly valued oysters.
Romans fished and imported them from all over European and Mediterranean coastlines and
extensively cultivated them (Günther 1897), and in some British estuaries there are archaeological
signs of overexploitation of native oyster beds since the first century (Rippon 2000). For centuries,
Ostrea edulis reefs supported a productive commercial fishery (Mackenzie et al. 1997). In the
eighteenth and nineteenth centuries, large offshore oyster grounds in the southern North Sea and
the English Channel produced up to 100 times more than today’s 100–200 t (U.K. Biodiversity
Group 1999, Berghahn & Ruth 2005). The richest natural oyster beds in Europe until the nineteenth
century were probably around Britain, from Stornoway to the Solway in the west and from the
Orkney Islands to the Firth of Forth in the east (Berghahn & Ruth 2005). In the mid-nineteenth
century these were heavily exploited; dredging of the oyster beds was one of the largest fisheries,
employing about 120,000 men around the coast in the 1880s (Tyler-Walters 2001), with an annual
yield of >50 million oysters (Berghahn & Ruth 2005). Oyster reefs at Strangford Lough, in Ireland,
once supported up to 20 boats employed in oyster dredging (Kennedy & Roberts 1999). In the
Wadden Sea, the commercial fishery for oysters started in the eleventh century and flourished in
the eighteenth century (Figure 13): in 1765 large oyster beds between Texel and Wieringen sup-
ported profitable fishery by 145 vessels, with catches over 100,000 oysters yr−1 vessel−1 (Wolff 2005).
By the late nineteenth century, beds of O. edulis were already severely depleted or physically
destroyed around most European coasts (Ocean Studies Board 2004). Regulations and fishery
closures were imposed in some regions. In the Wadden Sea, for example, management strategies
including minimum landing size, fishery closures, a licence system and maximum yield per year
have been applied since the seventeenth century (Berghahn & Ruth 2005). Similar initiatives were
taken in France. The decline could not, however, be halted and in the twentieth century catches
collapsed (e.g., Figure 13A,B). Overfishing and wasteful exploitation, combined with outbreaks of
diseases, habitat loss and change or destruction, reduction in water quality and other large-scale
environmental alterations, adverse weather conditions, and the introduction of non-native oysters
(and associated parasites and diseases, such as the protozoan Bonamia ostreae) for aquaculture and
other non-native species (e.g., the invasive gastropod Crepidula fornicata) were blamed for the
decline (Korringa 1952, Mackenzie et al. 1997, Wolff 2000, Jackson 2003, Berghahn & Ruth 2005).
Virtual extinction of native oyster beds has been documented in the Wadden Sea, where wild oysters
largely disappeared by 1950 (Wolff 2000, 2005, Lotze 2005); in Helgoland (Germany), where beds
largely disappeared by the mid-1900s (Korringa 1952, Franke & Gutow 2004, OSPAR Commission
2005); in the Dutch Easter Scheldt (van den Berg et al. 2005); in Belgium (OSPAR Commission
381
0.3 A
0.2
0.1
0.0
1770 1780 1790 1800 1810 1820 1830
6
Oysters harvested (× 106)
B
5
4
3
2
1
0
1870 1880 1890 1900 1910 1920 1930
25
C
20
15
10
0
1970 1975 1980 1985 1990 1995 2000
Figure 13 Annual landings of native European oysters, Ostrea edulis, (A) in the east Frisian Wadden Sea
during 1770–1830, (B) in the north Frisian Wadden Sea during 1868–1930 and (C) landings of cultured Pacific
oysters, Crassostrea gigas, in the Netherlands during 1970–2000. (From Lotze 2005. With permission.)
2005); in all deeper waters of the southern North Sea, such as in the Oyster Grounds (OSPAR
Commission 2005); in most areas of Galicia (Cano & Rocamora 1996) and in some bays in the
Black Sea (Zaitsev 2006). In the Firth of Forth (Scotland), which in past centuries had hosted one of
the most famous oyster banks, no oysters were found in 1957 (Dodd 2005). Dramatic stock decreases
have been reported as well on the Atlantic coasts in French Brittany, the Netherlands, Denmark,
Norway, Ireland and England and in the Mediterranean Sea (Korringa 1952, Mackenzie et al. 1997,
U.K. Biodiversity Group 1999, Barnabe & Doumenge 2001). In the United Kingdom, where 700
million oysters were consumed in London alone in 1864, the catch fell from 40 million in 1920
to 3 million in the 1960s and has not recovered (Tyler-Walters 2001). In Archachon Basin (France),
382
wild oysters, which had been exploited for ages, were nearly commercially exhausted in the mid-
1800s, leading to the introduction first of the Portuguese hollow oyster (Crassostrea angulata) and
then of the Pacific oyster (Crassostrea gigas).
Trends and threats
There is limited information about current trends and threats to remaining native oyster reefs in
Europe. Ostrea edulis is a relatively long-lived species and reproduces sporadically (Korringa 1952).
Thus, presumably, times of recovery from overexploitation or other causes of damage are very long
and are estimated to take up to 20 yr (Jackson 2003). O. edulis is considered to be highly sensitive
to substratum loss, smothering, contamination by synthetic compounds (particularly tributyltin
(TBT) antifouling paints used on ships and leisure craft, which, in the early 1980s, caused stunted
growth of oysters and probably affected reproductive capacity), oxygen depletion, reduced fresh-
water inputs, introduction of microbial pathogens/parasites, introduction of non-native species and
direct extraction (U.K. Biodiversity Group 1999, Jackson 2003, Hiscock et al. 2005). All these
factors impair recovery as well as restoration efforts. The main factors that probably threaten native
oyster reefs nowadays include illegal fishing as well as by-catch in trawling targeting other species,
poor water quality and pollution, changes to the environment (e.g., habitat loss due to coastal
development) and the introduction of non-native competitors, predators and diseases (Jackson 2003,
OSPAR Commission 2005).
Protection measures
Nowadays, the sparse remains of wild native oyster beds are probably one of the most endangered
marine habitats in Europe. Ostrea edulis, however, does not seem to be the target of any specific
protection measure, conservation legislation or convention at a European level. ‘Reefs’, including
biogenic reefs, are listed as a conservation feature in Annex I of the Habitats Directive; however,
native oyster reefs are not mentioned as a component (EC 2003). Since 2003, O. edulis beds are
included in the OSPAR list of threatened and/or declining species and habitats for all OSPAR areas
(OSPAR Commission 2005). Ostrea edulis is also included in the ‘Red’ lists of some regions (e.g.,
Wadden Sea, Black Sea). Indirect protection to native oyster reefs may also come from a number
of EC Directives related to shellfish, such as the 95/70/EC, which sets community-wide rules to
prevent the introduction and spread of the most serious diseases affecting bivalve molluscs, and
the Shellfish Waters Directive (Table 1).
Fisheries for native oysters are regulated (sometimes prohibited) at a national level (e.g.,
Hiscock et al. 2005, Zaitsev 2006) but other national or regional conservation initiatives seem to
be rare. There is little evidence that this management is leading to recovery of stocks. In the United
Kingdom, O. edulis is included in a Species Action Plan under the U.K. Biodiversity Action Plan
(U.K. Biodiversity Group 1999) and naturally occurring native oyster beds are considered a nation-
ally scarce habitat (Jackson 2003), although complex regulations still allow some harvesting.
Biogenic habitats: maerls

Maerls (also known as rhodolith beds) comprise several species of crust-forming, free-living (i.e.,
unattached), calcareous red algae (Donnan & Moore 2003). Over time, they can become abundant
enough to form substantial banks of live and dead material, with some European beds dated as
older than 5500 yr (Grall & Hall-Spencer 2003). The major maerl-forming species in European
waters are Phymatolithon calcareum, Lithothamnion corrallioides and L. glaciale. They can occur
383
in exposed and sheltered environments, from the surface down to 100 m in depth (e.g., near Corsica
and Malta), but most typically occur at 20–30 m (OSPAR Commission 2005).
Maerl beds are structurally and functionally complex habitats that support rich and diverse
assemblages and host many species unique to those habitats. There is also growing evidence that
maerl beds have considerable value as nursery grounds for species of commercial interest (Barbera
et al. 2003).
Records of the presence or absence of maerl biotopes on European coasts are patchy (Birkett
et al. 1998a). Detailed studies of maerl habitats have been undertaken only in the past 40 yr and at
only a few locations, mainly in France, Norway and Ireland. Large, historically accessible maerl banks
are relatively well recorded as a result of commercial interests. The locations of other maerl sites are
known from the results of grab-and-dredge sampling during scientific research cruises. In more recent
times, scuba divers have reported maerl banks. However, the extent of a maerl bed at any given
location, its species composition and the species associated with it remain largely unknown (Birkett
et al. 1998a). The most recent and comprehensive overview of maerl beds in Europe was compiled
under the EC MAST-funded BIOMAERL project (Donnan & Moore 2003). Unpublished databases
are expanding for the United Kingdom and France but an overall inventory has not been attempted.
In Europe maerl beds are patchily distributed and relatively restricted in size (Donnan & Moore
2003). They are found throughout the Mediterranean Sea, with important beds in Algeria, at
Marseilles, in Corsica and Sardinia and in the Aegean (Birkett et al. 1998a). Maerl beds are also
common on the Atlantic coasts, from Norway to Portugal. Spanish maerl deposits are confined
mainly to the Ria de Vigo and Ria de Arosa (Galicia, northwest Spain). Maerl beds are relatively
rare in the eastern English Channel, Irish Sea, North Sea and Baltic Sea (Barbera et al. 2003),
whereas they are particularly abundant in Brittany, with more that 70 beds >1 km2 and some of
the largest and thickest beds in Europe and the world (Grall & Hall-Spencer 2003). In Ireland,
maerl is widely distributed in the south and southwest (e.g., Galway Bay, Bantry Bay, Roaringwater
Bay; De Grave & Whitaker 1999), and Scotland is home to some of the most extensive maerl beds
in Europe (Birkett et al. 1998a).
Information on the status of present maerl beds in Europe is limited. Most Breton maerl beds
are affected by human activities and the only pristine grounds remaining are small compared with
the extensive maerl beds that covered several square kilometres in the 1960s (Grall & Hall-Spencer
2003). In 1999, surveys at one of the largest maerl beds in Brittany (Glenan), which was covered
with living maerl until extraction started some 35 yr ago, showed that live maerl was rare over
most of this bank while species-poor assemblages on muddy bottoms prevailed (Grall & Hall-
Spencer 2003). Even maerl beds included in Breton NATURA 2000 sites are far from pristine and
many are severely degraded.
Information on historical losses of maerl beds in Europe as a consequence of human activities is

virtually absent. Maerl has been harvested on a small scale in Europe for thousands of years for
use in animal food additives, water filtration systems, acid lake and pond treatment, biological
denitrification, toxin elimination, surfacing garden paths and in the pharmaceutical, cosmetics,
medical and nuclear industries, but mostly as a cost-effective source of calcium/magnesium soil
additive in agriculture and horticulture (Barbera et al. 2003, Grall & Hall-Spencer 2003). Initially,
the quantities extracted were small, being dug by hand from intertidal banks, but in the 1970s about
600,000 t of maerl were extracted per year in France alone (Birkett et al. 1998a). Maerl extraction
still forms a major part of the French seaweed industry, both in terms of tonnage and value of
harvest, although amounts have declined to about 500,000 t yr−1 (Grall & Hall-Spencer 2003). In
the United Kingdom and Ireland maerl has been harvested since the seventeenth century (De Grave
384
& Whitaker 1999) with up to 30,000 t yr−1of maerl harvested commercially in the River Fal from
1975 to 1991 (Birkett et al. 1998a). Currently only limited extraction of maerl takes place in the
United Kingdom and Ireland (De Grave & Whitaker 1999).
The extent to which this historical extraction has affected maerl beds is not known. Comparisons
between museum collections made in 1885–1891 and again in 1995–1997 at maerl beds in the
Firth of Clyde (Scotland) showed extensive changes, with substantial reduction in size and number
of living thalli of Phymatolithon calcareum (Hall-Spencer & Moore 2000). Such changes have
been attributed to mechanical impacts of scallop dredging, which started in that area in the 1930s
and became particularly intensive in the 1960s through the advent of more powerful boats, more
efficient dredges and better processing facilities. The wholesale removal of maerl habitats and
significant reductions in diversity and abundance to adjacent areas at five sites around the coasts
of Brittany have also been reported (Barbera et al. 2003) and attributed to commercial extraction
(Grall & Hall-Spencer 2003).
Trends and threats

Maerl habitats are considered highly sensitive to overexploitation and other human activities that
result in physical disturbance or deterioration in water quality (Barbera et al. 2003), particularly
smothering by fine sediments (Wilson et al. 2004). This sensitivity is compounded by long recovery
times due to the slow growth (approximately 1 mm yr−1) and accumulation characteristics of maerl
beds. The coralline algae that form the maerl are among the slowest-growing species, and substantial
deposits take centuries to millennia to accumulate (Hall-Spencer et al. 2003) so that any effects of
habitat removal are irreversible over timescales relevant to humans.
The major threats to maerl habitats have been recently reviewed (Barbera et al. 2003). The
most obvious threats are from the ongoing commercial extraction. The three main areas of com-
mercial exploitation in Europe have been Brittany, Cornwall and the west of Ireland. Nowadays,
only limited extraction takes place in Ireland and the United Kingdom (De Grave & Whitaker 1999)
but maerl extraction still forms a major part of the French seaweed industry (Grall & Hall-Spencer
2003). It has been predicted that if extraction rates persist at current levels the large Glenan deposit
(Britttany) could be exhausted within 50–100 yr (Grall & Hall-Spencer 2003).
In addition to the direct effects of harvesting, other direct and indirect impacts on maerl beds
have been noted. Damage to the surface of the beds is caused by towed demersal fishing gear, such
as scallop dredges, which significantly reduce bed complexity, biodiversity and long-term viability
(Hall-Spencer & Moore 2000, Barbera et al. 2003, Hall-Spencer et al. 2003). Hall-Spencer & Moore
(2000) found that scallop (Pecten maximus) dredging in the Clyde Sea led to a 70% reduction of
live maerl on a previously unexploited bed with no signs of recovery over the subsequent 4 yr.
Permanent moorings for pleasure boats can have similar, more localised, effects (Birkett et al.
1998a). The negative effects of increased eutrophication and turbidity in coastal waters both from
silt loads and nutrient runoff from agricultural land and aquaculture have been documented in
Galicia and in the Bay of Brest (Birkett et al. 1998a, Barbera et al. 2003). Smothering of maerl
beds as a consequence of the invasion of the gastropod Crepidula fornicata has been observed in
Breton bays (Grall & Hall-Spencer 2003). Maerl beds are also threatened by land reclamation and
proliferation of coastal structures that alter circulation patterns (Birkett et al. 1998a, Barbera et al.
2003).
Protection measures
Although maerl is confined to a relatively small proportion of European shallow sublittoral waters,
their conservation importance is being increasingly recognised (Birkett et al. 1998a, Donnan &
385
Moore 2003). Two of the most common maerl-forming species, Phymatolithon calcareum and
Lithothamnion corallioides, are now the only algal species specified as requiring appropriate
management measures under the Habitats Directive (Annex V). Free-living Corallinaceae are also
a named component of the habitat ‘Sandbanks which are slightly covered by sea water all of the
time’ (EC 2003). Maerl beds are also included in the OSPAR list and Mediterranean Red Book of
threatened habitats (Boudouresque et al. 1990, OSPAR Commission 2005).
In the United Kingdom, maerl is the subject of a Habitat Action Plan (U.K. Biodiversity Group
1999) and both L. corallioides and Phymatolithon calcareum are on the long list of species in the
U.K. Biodiversity Steering Group Report (U.K. Steering Group 1995). Furthermore, in the JNCC
interpretation of the EC Habitats Directive, maerl is identified as a key habitat within the Annex I
category ‘Sand banks which are slightly covered by seawater at all times’. This means that a number
of SACs being designated under the directive will provide protection to the maerl that they contain.
Maerl beds occur in three of 12 demonstration SACs within the United Kingdom, while the Fal
and Helford (Cornwall) candidate SAC includes the largest maerl bed in England. Recently, the
Board of Falmouth Harbour Commissioners in Cornwall has decided to cease licensing maerl
extraction (Hall-Spencer 2005).
France has also recognised biogenic reefs, such as maerls, as vulnerable habitats, and some
maerl grounds in Brittany lie within SACs (Grall & Hall-Spencer 2003). However, many of these
are already severely degraded and are affected by dredge fishing, eutrophication and the spread of
Crepidula fornicata. Maerl extraction in Brittany is under the control of the French mining man-
agement scheme, with quota schemes (80,000 t in 2001 on the Glenan bank) and regular environ-
mental surveys. However, such quotas are considered not compatible with regeneration of the
resource (Grall & Hall-Spencer 2003).
Biogenic formations
There are other examples of biogenic habitats that are severely affected and presumably have been
subject to massive losses over the centuries. However, most of these losses have probably passed
unnoticed and the information is scattered and mainly anecdotal. For example, off-shore rocky
formations in the north Adriatic Sea, both organogenic and fossil in nature, have been flattened
and reduced in size by trawling or other destructive forms of fisheries (Bombace 2001), sometimes
to virtual extinction. Mediterranean ‘coralligenous’ reefs, which are considered one of the most
valuable and diverse habitats in the Mediterranean Sea, are degraded and highly threatened by a
variety of human activities (Ballesteros 2006) but how much coralligenous habitat might have been
lost in the past as a consequence of these activities is not known. Significant declines in the extent
of wild intertidal mussel beds have been reported from large coastal areas of Germany, the Nether-
lands and Denmark, and Mytilus edulis beds are now rare in the Wadden Sea (OSPAR Commission
2005, Wolff 2005) and are considered under threat in the United Kingdom (Hiscock et al. 2005).
Large subtidal Sabellaria spinulosa reefs in the German Wadden Sea, which provided an important
habitat for a wide range of associated species, have been completely lost since the 1920s (U.K.
Biodiversity Group 1999, Wolff 2000) and similar losses have been reported also from areas of the
northeast Atlantic and the United Kingdom (OSPAR Commission 2005). A significant contraction
in the range of S. alveolata reefs on the south coast of England has occurred over a period of at
least 20 yr until 1984. Declines have also been reported in the western part of the north Cornish
coast, the upper parts of the Bristol Channel and in North Wales and the Dee Estuary but the causes
of this regression are not known (U.K. Biodiversity Group 1999).
The scarcity of information probably underlies the lack of adequate policies and protection
measures for these biogenic habitats. Biogenic reefs and concretions are all broadly covered as
‘Reef’ by the Habitats Directive but most are not specifically mentioned (EC 2003) and even
386
national and regional initiatives are limited. Coralligenous assemblages are listed in the Red Book
of Mediterranean assemblages (Boudouresque et al. 1990) and S. spinulosa reefs are included in
the Red List of Macrofaunal Benthic Invertebrates of the Wadden Sea and the OSPAR list of
threatened habitats (OSPAR Commission 2005). In the United Kingdom, both S. spinulosa and
S. alveolata are the subject of Habitat Action Plans (U.K. Biodiversity Group 1999).
Sedimentary habitats (mudflats, sandflats and subtidal soft bottoms)

Coastal areas are dominated by soft-sediment habitats. The marine biotope classification for Britain
and Ireland identifies a number of littoral and sublittoral sediment biotopes (U.K. Biodiversity
Group 1999). These are grouped into several major categories (gravels and sands, muddy sands,
muds and mixed sediments) and subdivided further according to depth (littoral, infralittoral or
circalittoral) and sediment size. Muddy habitats usually occur in sheltered areas, such as sea lochs,
enclosed bays and estuaries, whereas sandflats and coarser sediments tend to develop in more
exposed situations on the open coast. Distinctions are also sometimes made between estuarine and
marine habitats (e.g., OSPAR Commission 2005).
Despite the importance of these highly productive soft-sediment habitats, which support large
numbers of predatory birds and fishes, providing nursery, feeding and resting areas, and the long
history of studies on many aspects of their ecology, no comprehensive inventory of their extent
and status is available at a European level, and even regional or local initiatives are rare. The only
habitat for which there are some rough estimates of the amount and distribution are intertidal
mudflats. In the OSPAR region, the largest continuous area of intertidal mudflats borders the north
coasts of Denmark, Germany and the Netherlands in the Wadden Sea, covering around 4990 km2
(OSPAR Commission 2005). In the United Kingdom, intertidal mudflats are widespread, with
significant examples in the Wash, the Solway Firth, Mersey Estuary, Bridgwater Bay and Strangford
Lough; overall they are estimated to cover about 2700 km2 (U.K. Biodiversity Group 1999).
The extent of historical losses of soft-bottom habitats is virtually unknown for any country and
even regional or local information is scarce. Available information suggests that loss or deep
alteration of these types of habitats may have been extremely high, particularly in estuaries and
enclosed bays but there is also the possibility that unvegetated soft bottoms have increased their
area at the expense of losses in other more structurally complex habitats (e.g., seagrass beds). Past
losses are likely to have been related mainly to land claim for agriculture, ports and industrial and
urban developments. In the United Kingdom, for example, it is estimated that at least 88% of
estuaries have lost intertidal habitats and about 25% of overall estuarine intertidal flats have been
removed with peaks of up to 80% in some estuaries such as the Tees (U.K. Biodiversity Group
1999, OSPAR Commission 2005). The amount of intertidal mudflats and sandflats may have also
changed significantly over time in relation to coastal erosion, changes in sea levels and human
interventions to control these factors (e.g., Lee 2001).
Most often, however, these habitats have probably been deeply altered in their fundamental
characteristics, including sediment structure and composition, accretion or erosion rates, and inhab-
iting fauna. As an example, in Europe the massive use of hard defence structures and beach
replenishment schemes has deeply changed the structure of shallow surf-zone sediments along
whole coastlines in past decades (Airoldi et al. 2005, Martin et al. 2005), as is the case of the north
Adriatic Sea (e.g., Figure 5), presumably affecting an enormous and overlooked amount of shallow
387
soft-bottom habitats. Similarly, most sedimentary benthic systems on the continental shelf of Europe
have been modified by fishing activities, particularly bottom trawls and dredging, in the last 100 yr
(Ball et al. 2000, Frid et al. 2000). In the southern North Sea fishing is thought to have long been
the main ecological structuring force on the benthos, to an extent that makes it difficult even to
design robust field experiments due to the virtual lack of control areas (Hall-Spencer & Moore
2000). How much of this transformation should be considered as habitat loss or degradation is
difficult to quantify.
Trends and threats
There is limited organised information on the current trends and threats to sedimentary environments
(Brown & Mclachlan 2002). Today the major threats to sedimentary habitats are more likely to be
linked to further land claim, construction of marinas and slip ways, the widening and dredging of
channels for navigation, pipe and cable laying, oil and gas extraction, tourist developments and
infrastructures and the construction of sea defences. Some of these threats have slowed considerably
in recent years, at least in some European countries, but they have not stopped. In the United
Kingdom, many coastal areas, including estuaries, are now either licensed or available for explo-
ration and development (U.K. Biodiversity Group 1999). Pollution from sewage discharge, aquac-
ulture activities, industries and shipping are also important threats to sedimentary environments
and associated fauna, leading to anoxic conditions particularly in estuaries and enclosed basins, as
observed in Scandinavian and Baltic waters (Karlson et al. 2002), and to long-term accumulation
of contaminants (Islam & Tanaka 2004).
Physical disturbance by fishing and aggregate dredging activities also represents a major threat
to Europe’s sedimentary habitats and associated biota (Lindeboom & de Groot 1998, Tudela 2004),
although nowadays highly disturbed seabeds may appear to be relatively unaffected by fishing
activities or other physical disturbances (e.g., Hall-Spencer & Moore 2000). On the Dutch conti-
nental shelf, the fisheries are now so intensive that every square metre is trawled, on an average,
once to twice a year (Lindeboom 1995), and this broadly applies to the entire sea bed of the North
Sea (Gray 1997).
Direct extraction of sands and gravels for coastal developments, use in the construction industry
and beach nourishments are also major, increasing threats for sedimentary habitats (Newell et al.
1998, van Dalfsen et al. 2000). Extraction of sands has steadily increased in most north European
countries during the past few decades (ICES 2006b). In the Netherlands, for example, extraction
of sands has increased from <5 million m3 yr−1 in 1974 to >35 million m3 yr−1 in 2001, and in the
coming decades an average request of 19–43 million m3 yr−1 is expected. Much of the extracted
sand is used for beach recharges and coastal defence. Beach nourishments are being increasingly
used along European coasts as a ‘soft’ measure to counteract erosion (Hamm et al. 2002), but the
consequences of both the extraction and the disposal of sands on sedimentary habitats and biota
have received limited attention (Desprez 2000, van Dalfsen et al. 2000, Simonini et al. 2005).
Some projections of loss are available for intertidal areas in relation to possible future changes
in sea level, recession of coastlines and coastal ‘squeeze’. For example, sea-level rise is projected
to cause a loss of 80–100 km2 of intertidal flats in England between 1993 and 2013 (U.K.
Biodiversity Group 1999), particularly in southern and southeast regions; the major firths in Scotland
will probably also be affected.
Protection measures
Protection for intertidal and shallow mudflats and sandflats is provided by various international
and E.U. agreements, including the Ramsar Convention, the Bonn Convention, the Bern Convention,
388
and the Birds and Habitats Directives (Table 1). In particular, ‘Mudflats and sandflats not covered
by seawater at low tide’ and ‘Sandbanks which are slightly covered by sea water all the time’ are
listed in Annex I of the Habitats Directive. Mudflats are also included within several other designated
Annex I Habitats: ‘Estuaries’, ‘Lagoons’ and ‘Large shallow inlets and bays’. Mudflats are also in
the list of OSPAR threatened habitats (OSPAR Commission 2005). Some countries also have
national protection measures. For example, in the United Kingdom mudflats are the subject of a
Habitat Action Plan (U.K. Biodiversity Group 1999); furthermore, over 300 SSSIs including
mudflats have been designated in estuaries and 10 coastal ASSIs in Northern Ireland contain
significant areas of mudflats.
Soft bottoms deeper than 30 m do not seem to be the target of any specific protection measure
at a European level (Hiscock et al. 2005), although a number of EC Directives that regulate water
quality provide indirect protection from some types of impacts (Table 1). There are, however,
national initiatives. For example, in the United Kingdom ‘Sublitoral sands and gravels’ and ‘Mud
habitats in deep waters’ are the subjects of Habitat Action Plans (U.K. Biodiversity Group 1999).
Commercial fishing activities are excluded from a number of estuaries and bays around the coast
of the United Kingdom, which are important nursery areas for juvenile commercial species (e.g.,
River Exe, River Conwy and Filey Bay). Fishing activities are prohibited within 500 m of gas and
oil platforms, from firing ranges and in close proximity to certain military installations (U.K.
Biodiversity Group 1999).
Discussion
Population density along European coasts has been growing since ancient times. There are increas-
ingly greater demands and impacts on the habitats and resources in coastal environments. The
present review has shown that such intensive exploitation has caused dramatic losses and severe
deterioration of native coastal habitats (e.g., Tables 2–4). There are many policies and directives
Table 4 Summary of main characteristics of European coastlines and habitats based on

reviewed sources
Characteristic Value Main references
Coastline length a 325,892 km Pruett & Cimino 2000
Population within 50 kmb 200 × 106 Stanners & Bourdeau 1995
Degraded coastlines 85% EEA 1999a
Years of impactc 2500 yr Rippon 2006, Lotze et al. 2006
Artificial coastlines 22,000 km2 EEA 2005
Defended/eroding coastlines 7,600/20,000 km EC 2004
Increase in N/P loads 1940s–1980s 2- to 4-/4- to 8-fold Nehring 1992, EEA 2001, Karlson et al.
2002
Number of invasive species 450–600 Reise et al. 2006
MPAs (Number/Total surface) 1,129/ 236,000 km2 UNEP/WCMC 2006, MPA Global 2006
Present coastal wetlands/loss since 1900s 51,910 km2/>65% Nivet & Frazier 2004, EEA 2006a
Present seagrasses/historical lossesd 7290 km2/>65% Duarte 2002, Green & Short 2003
Present wild native oyster reefs/historical lossesd Scarce/>90% Mackenzie et al. 1997
Present macroalgal beds/historical lossesd Unknown/2–4 m in depth Vogt & Schramm 1991, Eriksson 2002
a Including islands.
b In the 1990s.
c Since beginning of modification and transformation of coastal landscapes.
d Estimate based on reviewed local to regional sources.
389
Table 5 Past (earlier than 1900 = P), recent (twentieth century = R) and present (N) main
drivers of habitat loss along European coasts based on reviewed sources
Wetlands Seagrasses Macroalgae Biogenic habitats Sediments
Impact P R N P R N P R N P R N P R N
Claim/conversion *** *** ** ** ** * * * * *** *** **
Coastal development *** *** *** *** *** *** ? ** ** *** *** ***
Coastal defence ** *** * ** g *ga ** ***
Exploitation ** * * *** *** *** * **
Water quality ** ** ** ** *** *** ? *** *** ** ** **
Diseases/pests/predators * * *** ** ? ** ** ** *** ***
Destructive fishing * *** * ? ** * ** *** *** ?b ?b
Aquaculture * ** * ** * ** ***
Note: Habitats are coastal wetlands (including salt marshes); seagrass meadows; macroalgal beds (kelps, fucoids and
other complex macroalgae); biogenic habitats (including oyster reefs and maerls); and sedimentary habitats (mudflats,
sandflats and subtidal soft bottoms). Impacts are drainage, embankment, land claim and habitat conversion (e.g., into
agriculture land or into freshwater lake); coastal development including urban and industrial developments, ports and
infrastructures, marinas and tourist and recreational developments; coastal defence which includes hard structures, beach
nourishments and other measures associated with impacts from erosion, storms and sea-level rise; direct exploitation or
harvesting; water quality including organic and chemical pollution and altered sedimentation regimes; outbreaks of
diseases, introduced species, competitors or predators; destructive fishing techniques (e.g., trawling); aquaculture. Blank
cells = nil or modest, * = low, ** = moderate/locally high, *** = high and widespread, g = habitat gain, ? = not known.
a Negative effects of beach nourishments, enhancement of substratum availability from hard structures.
b Fishing affects soft bottoms extensively, but it is difficult to evaluate how many soft sediment habitats are lost.
aimed at reducing and reversing these losses (e.g., Table 1) but their overall positive benefits have
been low.
Coastal habitats are affected by many threats with a few dominant threats that vary somewhat
by habitat and over time (Table 5). The greatest impacts to wetlands have been land claim and
coastal development with the latter rising in importance over time. The greatest impacts to sea-
grasses and macroalgae are presently associated with degraded water quality whereas in the past
there have been more effects from destructive fishing and diseases. Coastal development remains
an important threat to seagrasses in particular. For biogenic habitats, some of the greatest impacts
have been from destructive fishing and overexploitation with additional impacts of disease, partic-
ularly to native oysters. Coastal development and defence have had the greatest known impacts on
soft-sediment habitats with a high likelihood that trawling has impacted vast areas while not causing
loss of soft-sediment habitats per se.
Shellfish, oyster reefs in particular, have been among the most severely affected of all coastal
habitats by overexploitation and other human-driven changes to the environment. By the late
nineteenth century, overfishing combined with outbreaks of diseases, habitat transformation, and
the introduction of non-native competitors and parasites, had already wiped out wild Ostrea edulis
reefs around much of the European coastline. Where documentation is available, it is evident that
the loss of shellfish habitat is a major cause of species extirpation and declines in biodiversity
(Wolff 2000, Lotze et al. 2005). Currently most native oyster reefs are functionally or entirely
extinct in most coastal areas of Europe and they are probably one of the most endangered marine
habitats. However, loss and threats to these habitats are largely overlooked and shellfish beds do
not seem to be covered by adequate protection measures, conservation legislation or convention at
a European level. Oyster reefs are not mentioned specifically in the Habitats Directive, which sets
the present framework for habitat protection in Europe. Although less documented and more
390
localised, similar losses seem to have occurred also to other types of shellfish reefs and biogenic
formations, such as maerl beds, intertidal mussels beds, Sabellaria spp. reefs and coralligenous
formations.
The current losses in European coastal habitats are alarming; even worse is the fact that these
losses are only measured against recent distributions with little recognition of the compounding
impact of centuries and millennia of habitat loss. This historical short-sightedness has been referred
to as ‘shifting baselines’, where we only recognise declines in the natural environment relative to
the baseline of recent memory in each generation (sensu Dayton et al. 1998, Jackson et al. 2001).
At present, there seems to be limited public, political and even scientific awareness of the extent,
importance and consequences of such a long history of coastal habitat loss (Lotze 2004).
The evidence reviewed in the present work clearly indicates that in some European regions
most estuarine and near-shore coastal habitats were probably already severely degraded or driven
to virtual extinction well before 1900 and sometimes much earlier than that. Ecological descriptions
of coastal marine habitats are rather recent (mid-1900s), and long-term documentation of habitat
declines and losses are virtually missing for most systems. Even when documentation is available,
evidence tends to be overlooked. For example, Wolff (2000) points out how most Dutch people are
inclined to ignore the profound habitat changes and losses that occurred during the past 2000 yr
in the Wadden Sea, and they tend to consider present-day systems to be in a ‘natural’ state.
Nowadays only a small percentage of the European coastline is considered in ‘good’ condition
(EEA 1999c). The degradation has continued at high rates over the past few decades. In France,
for example, it is estimated that 15% of natural areas on the coast have disappeared since 1976
and are continuing to do so at the rate of 1% a year (Stanners & Bourdeau 1995). In Italy, around
7000 km2 of coastal marshes were present at the beginning of the 1900s, no more than 1920 km2
in 1972 and fewer than 1000 km2 today (Stanners & Bourdeau 1995). Losses of coastal wetlands
and seagrasses exceeding 50% of original area have been documented for most countries where
long-term data were available, with peaks above 80% for many regions. Beds of complex macro-
algae have been under severe recession since at least the early 1900s. An impressive number of
local to regional extinctions of habitats have been documented. Overall, it is estimated that every
day between 1960 and 1995, a kilometre of ‘unspoilt’ European coastline has been developed
(EUCC 1998). Those fragments of native habitats that remain are under continued threat.
Recommendations for conservation and management

Fortunately there is recognition at the European level that the current management and conservation
of marine diversity and habitats is insufficient. In particular, there have been recent and significant
E.U. policies to identify the coastal and marine habitats of Europe and to develop networks of
MPAs around them. These efforts have been inspired in part by the international commitments
developed as part of the Rio CBD and the World Summit for Sustainable Development. The
development of E.U. MPAs is included in the Habitats Directive and the Birds Directive and
embodied in particular in the development of protected areas in the NATURA 2000 network. There
are also other regional (e.g., OSPAR) and national initiatives with similar aims for protection of
endangered habitats and species (Table 1). These policies are necessary but not sufficient. Indeed
MPAs alone have limited use for addressing threats such as poor water quality, disease or even
coastal development. Some key needs and opportunities for enhancing the overall conservation and
management of coastal and marine habitats in Europe are suggested below.
Currently, there is no comprehensive summary of the distribution of habitats along European
coastlines and their management is not well informed by adequate knowledge of their distribution
and status. Despite millennia of reliance on the resources from coastal and marine ecosystems, we
cannot accurately describe the distribution of even the shallowest habitats. Detailed habitat mapping
391
should therefore be given high priority to promote conservation and sustainable management
practices. Some databases are available or are being prepared for some countries or habitats but
most information is scattered, fragmented or limited to few case studies. At least there has been
good progress in developing a consistent habitat classification as part of the European Union Nature
Information System (EUNIS) (Davies et al. 2004), which is a necessary precursor to developing a
consistent database of habitat distribution.
These habitat distributions then need to be compared with the existing protected areas to identify
gaps in protection. To fill these gaps, there should be systematic planning for placement of new
protected areas and other management measures. Increasingly scientists, agencies and organisations
are using systematic planning approaches to target the placement of their protection and manage-
ment efforts particularly at regional levels (e.g., Beck & Odaya 2001, Possingham et al. 2002,
Airamè et al. 2003, Leslie et al. 2003, Beck et al. 2004). These approaches enable decision makers
to develop a range of solutions for protection and management and to examine how changes in
decisions can affect solutions.
The values of these coastal habitats also need to be better assessed to provide real estimates
of the ecosystem services that they provide such as pollution regulation, storm hazard reduction,
productivity of nurseries for fisheries and recreation (Agardy & Alder 2005). Better valuations of
these services will illustrate for communities and governments the real costs of this habitat loss
and should provide impetus and economic incentives for their protection and restoration.
There are still reasonable opportunities for conservation of coastal habitats in key areas through-
out Europe. This protection should be put in place quickly because conservation is cheaper than
restoration. The current EC policies are mostly aimed at these protections but they do need to be
implemented.
Given the extent of damage to coastal habitats, restoration will be required in many places to
meet any reasonable goals for conservation and management. Information on historical distributions
and loss is extremely important because management goals for these habitats should be based on
historical estimates of the distributions of these habitats, not the vastly reduced current distributions
(Beck 2003). Even extremely modest goals of 10% protection of the historical distributions of
European coastal habitats will require some restoration for many habitats.
To meet these goals for conservation and restoration there should be greater involvement by
nongovernmental organisations and community groups and there are tools that they can use to
contribute directly to conservation and management. These groups have often been involved in
efforts to develop MPAs and restore coastal habitats and these efforts should be encouraged and
expanded. There are also new tools, such as the private leasing and ownership of marine lands and
resources that can be employed more often by private groups to help protect and restore these
coastal habitats (Beck et al. 2004). For example, the National Trust in the United Kingdom leases
intertidal lands and sea beds from the government along some 180 km of the coast for conservation
and restoration. Recently there has been new policy adopted by the French government that allows
private groups also to lease subtidal lands as is commonly done by many business interests (e.g.,
aquaculture industry).
Most coastal habitats lie within the exclusive economic zones of individual countries and thus
the individual coastal zone management of these countries must be strengthened to protect and
manage these habitats. The European Union and member nations have been dedicated to developing
better Integrated Coastal Zone Management (ICZM) for some time. The development of strong
and effective ICZM programmes has been slow for most nations. These programmes need to
advance further to slow and reverse coastal habitat loss.
There is much academic and agency interest in developing a more ecosystem-based manage-
ment (E-BM) approach for managing the many marine resources and the overlapping stakeholder
needs for access to these resources (e.g., Browman & Stergiou 2005). E-BM has been incorporated
392
as a central goal of the European Union’s emerging ‘Marine Strategy’ (Table 1). While this approach
is needed and sensible, it will take many years to develop and its development should not be allowed
to slow efforts to protect and restore habitats now.
Conclusions
Europe has seen decades, centuries, even millennia of coastal habitat loss and it continues today.
Estuaries, enclosed bays and near-shore shelf habitats around the European coasts are some of the
most degraded environments on Earth as they have been used intensively for thousands of years.
These functionally valuable coastal ecosystems are still a focal point for human colonisation and
use. Before all the services from these ecosystems are lost, efforts should be redoubled to protect
the remaining habitats, slow and stop future losses and restore some of these habitats.
Recent efforts toward a more sustainable use of these coastal resources have not reversed the
trend. There is no single best strategy for addressing these losses and there are many approaches
that can help. Many of these are commonsense recommendations that are common in policies
within and between nations, but the progress toward them has been slow with a few exceptions.
The United Kingdom clearly has made more progress than most nations in the European Union
and internationally. If the long history of physical destruction, fragmentation and transformation
is further neglected, and if significant, irreversible thresholds are passed, the future sustainability
of those few fragments of native or semi-native habitats that remain may ultimately and finally be
compromised.
Acknowledgements
The work was supported by a grant from The Nature Conservancy. L.A. was further supported by
an Assegno di Ricerca of the University of Bologna. We are grateful to Heike Lotze and Keith
Hiscock for valuable input and for comments on an earlier draft of the work and to Robin Gibson
for his unfailing support. Many other people stimulated our work and gave useful input/references,
including Claudio Battelli, Erik Bonsdorff, Paolo Guidetti, Fiorenza Micheli, Karsten Reise and
Enric Sala. We are grateful to all the authors who provided figures from their work and the publishers
who agreed to their publication. We thank particularly Klemens Erikkson, who kindly prepared
original figures for this work, and Sandy Beck for lending her artistry to Figure 2. We also wish
to thank Giovanna Branca, Kendra Karr, Caitlyn Toropova, Dan Dorfman, Chris Shepard and Marco
Abbiati for their support at various stages of the work. L.A. is particularly grateful to Elena Fuschini
for her invaluable assistance with library searches. This publication is contribution number MPS-
07016 of the E.U. network of Excellence MarBEF.
References
Adam, P. 1990. Saltmarsh Ecology. Cambridge: Cambridge University Press.
Adam, P. 2002. Saltmarshes in a time of change. Environmental Conservation 29, 39–61.
Agardy, T. & Alder, J. 2005. Coastal systems. In Millennium Ecosystem Assessment: Strengthening Capacity
to Manage Ecosystems Sustainably for Human Well Being, Volume 1, Conditions and Terms. R. Hassan
et al. (eds). Washington, D.C.: Island Press, 513–549.
Airamè, S., Dugan, J.E., Lafferty, K.D., Leslie, H., McArdle, D.A. & Warner, R.R. 2003. Applying ecological
criteria to marine reserve design: a case study from the California Channel Islands. Ecological
Applications 13 (Suppl.), 170–184.
Airoldi, L. 1998. Roles of disturbance, sediment stress, and substratum retention on spatial dominance in algal
turf. Ecology 79, 2759–2770.
393
Airoldi, L. 2003. The effects of sedimentation on rocky coast assemblages. Oceanography and Marine Biology:
An Annual Review 41, 161–263.
Airoldi, L., Abbiati, M., Beck, M.W., Hawkins, S.J., Jonsson, P.R., Martin, D., Moschella, P.S., Sundelöf, A.,
Thompson, R.C. & Åberg, P. 2005. An ecological perspective on the deployment and design of low-
crested and other hard coastal defence structures. Coastal Engineering 52, 1073–1087.
Allen, H.D. 2003. A transient coastal wetland: from estuarine to supratidal conditions in less than 2000 years —
Boca do Rio, Algarve, Portugal. Land Degradation and Development 14, 265–283.
Allen, J.R.L. 1997. The geo-archaeology of land-claim in coastal wetlands: a sketch from Britain and the
north-west European Atlantic and North Sea coasts. Archaeological Journal 154, 1–54.
Allen, J.R.L. 2000. Morphodynamics of Holocene salt marshes: a review sketch from the Atlantic and southern
North Sea coasts of Europe. Quaternary Science Reviews 19, 1155–1231.
Allen, J.R.L. & Pye, K. (eds) 1992. Saltmarshes: Morphodynamics, Conservation and Engineering Signifi-
cance. Cambridge: Cambridge University Press.
Alongi, D.M. 2002. Present state and future of the world’s mangrove forests. Environmental Conservation
29, 331–349.
Anthony, E.J. 1994. Natural and artificial shores of the French Riviera. An analysis of their interrelationships.
Journal of Coastal Research 10, 48–58.
Bacchiocchi, F. & Airoldi, L. 2003. Distribution and dynamics of epibiota on hard structures for coastal
protection. Estuarine Coastal and Shelf Science 56, 1157–1166.
Badalamenti, F., Ramos, A.A., Voultsiadou, E., Sánchez Lizaso, J.L., D’Anna, G., Pipitone, C., Mas, J., Ruiz
Fernandez, J.A., Whitmarsh, D. & Riggio, S. 2000. Cultural and socio-economic impacts of Medi-
terranean Marine Protected Areas. Environmental Conservation 27, 110–125.
Baden, S., Gullström, L.B., Pihl, L. & Rosenberg, R. 2003. Vanishing seagrass (Zostera marina, L.) in Swedish
coastal waters. Ambio 32, 374–377.
Ball, B.J., Fox, G. & Munday, B.W. 2000. Long- and short-term consequences of a Nephrops trawl fishery
on the benthos and environment of the Irish Sea. ICES Journal of Marine Science 57, 1315–1320.
Ballesteros, E. 1992. Els Vegetals i La Zonació Litoral: Especies, Comunitats i Factors Que Influeixem en la
Seva Distribució. Arxius Secció Ciéncies 101. Barcelona: Institut d’Estudis Catalans.
Ballesteros, E. 2006. Mediterranean coralligenous assemblages: a synthesis of present knowledge. Oceanog-
raphy and Marine Biology: An Annual Review 44, 123–195.
Balmford, A., Green, R.E. & Jenkins, M. 2003. Measuring the changing state of nature. Trends in Ecology
and Evolution 18, 326–330.
Barbera, C., Bordehore, C., Borg, J.A., Glémarec, M., Grall, J., Hall-Spencer, J.M., de la Huz, Ch., Lanfranco,
E., Lastra, M., Moore, P.G., Mora, J., Pita, M.E., Ramos-Esplá, A.A., Rizzo, M., Sánchez-Mata, A.,
Seva, A., Schembri, P.J. & Valle, C. 2003. Conservation and management of northeast Atlantic and
Mediterranean maerl beds. Aquatic Conservation: Marine and Freshwater Ecosystems 13 (Suppl.),
65–76.
Barmawidjaja, D., van der Zwaan, G., Jorissen, F. & Puskaric, S. 1995. One hundred fifty years of eutroph-
ication in the northern Adriatic Sea: evidence from a benthic foraminiferal record. Marine Geology
122, 367–384.
Barnabe, G. & Doumenge, F. 2001. Mariculture of Mediterranean species. In Encyclopaedia of Ocean Sciences.
Volume 3, J.H. Steele et al. (eds). San Diego, California: Academic Press, 1567–1571.
Bartsch, I. & Tittley, I. 2004. The rocky intertidal biotopes of Helgoland: present and past. Helgoland Marine
Research 58, 289–302.
Bavestrello, G., Cerrano, C., Zanzi, D. & Cattaneo-Vietti, R. 1997. Damage by fishing activities to the
gorgonian coral Paramuricea clavata in the Ligurian sea. Aquatic Conservation: Marine and Fresh-
water Ecosystems 7, 253–262.
Beck, M.W. 2003. The sea around: marine regional planning. In Drafting a Conservation Blueprint: A Practi-
tioners’ Guide to Planning for Biodiversity, C.R. Groves (ed.). Washington, D.C.: Island Press, 319–344.
Beck, M.W., Heck, K.L., Jr., Able, K.W., Childers, D.L., Eggleston, D.B., Gillanders, B.M., Halpern, B.,
Hays, C.G., Hoschino, K., Minello, T.J., Orth, R.J., Scheridan, P.F. & Weinstein, M.P. 2001. The
identification, conservation, and management of estuarine and marine nurseries for fish and inverte-
brates. BioScience 51, 633–641.
394
Beck, M.W., Marsh, T.D., Reisewitz, S.E. & Bortman, M.L. 2004. New tools for marine conservation: the
leasing and ownership of submerged lands. Conservation Biology 18, 1214–1223.
Beck, M.W. & Odaya, M. 2001. Ecoregional planning in marine environments: identifying priority sites for
conservation in the northern Gulf of Mexico. Aquatic Conservation-Marine and Freshwater Ecosys-
tems 11, 235–242.
Benedetti-Cecchi, L., Pannacciulli, F., Bulleri, F., Moschella, P.S., Airoldi, L., Relini, G. & Cinelli, F. 2001.
Predicting the consequences of anthropogenic disturbance: large-scale effects of loss of canopy algae
on rocky shores. Marine Ecology Progress Series 214, 137–150.
Berghahn, R. & Ruth, M. 2005. The disappearance of oysters from the Wadden Sea: a cautionary tale for no-
take zones. Aquatic Conservation: Marine and Freshwater Ecosystems 15, 91–104.
Bianchi, C.N. & Peirano, A. 1995. Atlante delle Fanerogame Marine della Liguria. Posidonia oceanica e
Cymodocea nodosa. La Spezia: ENEA CRAM, Segraf.
Bird, E.C.F. 1993. Submerging Coasts. The Effects of a Rising Sea Level on Coastal Environments. Chichester,
U.K.: John Wiley & Sons.
Birkett, D.A., Maggs, C.A. & Dring, M.J. 1998a. Maerl (Volume 5). An Overview of Dynamic and Sensitivity
Characteristics for Conservation Management of Marine SACs. Scottish Association for Marine
Science (U.K. Marine SACs Project). Online. Available HTTP: http://www.ukmarinesac.org.uk/pdfs/
maerl.pdf (accessed 2 August 2006).
Birkett, D.A., Maggs, C.A., Dring, M.J., Boaden, P.J.S. & Seed, R. 1998b. Infralittoral Reef Biotopes with
Kelp Species (Volume 7). An Overview of Dynamic and Sensitivity Characteristics for Conservation
Management of Marine SACs. Scottish Association for Marine Science (U.K. Marine SACs Project).
Online. Available HTTP: http://www.ukmarinesac.org.uk/pdfs/reefkelp.pdf (accessed 6 August 2006).
Bokn, T.L., Moy, F.E., Magnusson, J.B. & Murray, S.N. 1992. Changes in fucoid distributions and abundances
in the Inner Oslofjord, Norway: 1974–1980 versus 1988–1990. Acta Phytogeographica Suecica 78,
117–124.
Bombace, G. 2001. Influence of climatic changes on stocks, fish species and marine ecosystems in the
Mediterranean sea. Archivio di Oceanografia e Limnologia 22, 67–72.
Bondesan, M., Castiglioni, G.B., Elmi, C., Gabbbianelli, G., Marocco, R., Pirazzoli, P.A. & Tomasin, A. 1995.
Coastal areas at risk from storm surges and sea-level rise in northeastern Italy. Journal of Coastal
Research 11, 1354–1379.
Boorman, L.A. 2003. Saltmarsh Review. An Overview of Coastal Saltmarshes, Their Dynamic and Sensitivity
Characteristics for Conservation and Management. JNCC Report, No. 334. Peterborough, U.K.:
JNCC. Online. Available HTTP: http://www.jncc.gov.uk/page-2347 (accessed 27 July 2006).
Boström, C., Baden, S.P. & Krause-Jensen, D. 2003. The seagrasses of Scandinavia and the Baltic Sea. In
World Atlas of Seagrasses, E.P. Green & F.T. Short (eds). Berkeley: University of California Press,
UNEP-WCMC, 27–37.
Boudouresque, C.F., Meinesz, A., Ballesteros, E., Ben Maiz, N., Boisset, F., Cinelli, F., Cirik, S., Cormaci,
M., Jeudy de Grissac, A., Laborel, J., Lanfranco, E., Lundberg, B., Mayhoub, H., Panayotidis, O.,
Semroud, R., Sinnassamy, J.M. & Span, A. 1990. Livre Rouge “Gérard Vuignier” des Végétaux,
Peuplements et Paysages Marins Menacés de Méditerranée. Map Technical Report Series, 43. Athens:
UNEP/IUCN/GIS Posidonie.
Boudouresque, C.F. & Verlaque, M. 2002. Biological pollution in the Mediterranean Sea: invasive versus
introduced macrophytes. Marine Pollution Bulletin 44, 32–38.
Bray, M.J. & Hooke, J.M. 1997. Prediction of soft-cliff retreat with accelerating sea-level rise. Journal of
Brooks, T.M., Mittermeier, R.A., Mittermeier, C.G., da Fonseca, G.A.B., Rylands, A.B., Konstant, W.R., Flick,
P., Pilgrim, J., Oldfield, S., Magin, G. & Hilton-Taylors, C. 2002. Habitat loss and extinction in the
hotspots of biodiversity. Conservation Biology 16, 909–923.
Browman, H.I. & Stergiou, K.I. 2005. Introduction: politics and socio-economics of ecosystem-based man-
agement of marine resources. Marine Ecology Progress Series 300, 241 only.
Brown, A.C. & Mclachlan, A. 2002. Sandy shore ecosystems and the threats facing them: some predictions
for the year 2025. Environmental Conservation 29, 62–77.
395
Bulleri, F. & Airoldi, L. 2005. Artificial marine structures facilitate the spread of a non-indigenous green alga,
Codium fragile ssp. tomentosoides, in the north Adriatic Sea. Journal of Applied Ecology 42, 1063–1072.
Burke, L., Kura, Y., Kassem, K., Revenga, C., Spalding, M. & Mc Allister, D. 2001. Pilot Analysis of Global
Ecosystems. Coastal Ecosystems. Washington, D.C.: WRI. Online. Available HTTP: http://pdf.wri.org/
page_coastal.pdf (accessed 4 August 2006).
Cano, J. & Rocamora, J. 1996. Growth of the European flat oyster in the Mediterranean Sea (Murcia, SE
Spain). Aquaculture International 4, 67–84.
Caressa, S., Ceschia, C., Orel, G. & Treleani, R. 1995. Present and former populations in the Gulf of Trieste
between Punta Salvatore and Punta Tagliamento (upper Adriatic). In Posidonia Oceanica. A Contri-
bution to the Preservation of a Major Mediterranean Marine Ecosystem, F. Cinelli et al. (eds). Rivista
Maritima 12 (Suppl.), 174–187.
Ceccherelli, G. & Cinelli, F. 1997. Short-term effects of nutrient enrichment of the sediment and interactions
between the seagrass Cymodocea nodosa and the introduced green alga Caulerpa taxifolia in a
Mediterranean bay. Journal of Experimental Marine Biology and Ecology 217, 165–177.
Cencini, C. 1998. Physical processes and human activities in the evolution of the Po delta, Italy. Journal of
Christie, H., Fredriksen, S. & Rinde, E. 1998. Regrowth of kelp and colonization of epiphyte and fauna
community after kelp trawling at the coast of Norway. Hydrobiologia 375–376, 49–58.
Cicogna, F., Bavestrello, G. & Cattaneo-Vietti, R. (eds). 1999. Red Coral and Other Mediterranean Octocorals:
Biology and Protection. Rome: Ministero Politiche Agricole.
Connell, S.D. 2005. Assembly and maintenance of subtidal habitat heterogeneity: synergistic effects of light
penetration and sedimentation. Marine Ecology Progress Series 289, 53–61.
Connor, D.W., Allen, J.H., Golding, N., Howell, K.L., Lieberknecht, L.M., Northen, K.O. & Beker, J.B. 2004.
The Marine Habitat Classification for Britain and Ireland. Version 04.05. Peterborough: JNCC.
Online. Available HTTP: http://www.jncc.gov.uk/MarineHabitatClassification (accessed 8 June 2006).
Cooper, N.J., Cooper, T. & Burd, F. 2001. Twenty-five years of salt marsh erosion in Essex: implications for
coastal defence and nature conservation. Journal of Coastal Conservation 7, 31–40.
Cormaci, M. & Furnari, G. 1999. Changes of the benthic algal flora of the Tremiti islands (southern Adriatic)
Italy. Hydrobiologia 398/399, 75–79.
Costanza, R., d’Arge, R., de Groot, R., Farber, S., Grasso, M., Hannon, B., Limburg, K., Naeem, S., O’Neill,
R.V., Paruelo, J., Raskin, R.G., Sutton, P. & van den Belt, M. 1997. The value of the world’s ecosystem
services and natural capital. Nature 387, 253–260.
Curtis, T.G.F. & Skeffington, M.J.S. 1998. The salt marshes of Ireland: an inventory and account of their
geographical variation. Biology and Environment: Proceedings of the Royal Irish Academy 98B,
87–104.
Davidson, N.C., Laffoley, D.d., Doody, J.P., Way, L.S., Gordon, J., Key, R., Drake, C.M., Pienkowski, M.W.,
Mitchell, R. & Duff, K.L. 1991. Nature Conservation and Estuaries in Great Britain. Peterborough,
U.K.: Nature Conservancy Council.
Davies, C.E., Moss, D. & Hill, M.O. 2004. EUNIS Habitat Classification Revised 2004. Report to EEA and
European Topic Centre on Nature Protection and Biodiversity. October 2004. Online. Available HTTP:
http://eunis.eea.europa.eu/upload/EUNIS_2004_report.pdf (accessed 1 August 2006).
Davison, D.M. & Hughes, D.J. 1998. Zostera Biotopes (Volume 1). An Overview of Dynamics and Sensitivity
Characteristics for Conservation Management of Marine SACs. Scottish Association for Marine
Science (U.K. Marine SACs Project). Online. Available HTTP: http://www.ukmarinesac.org.uk/
zostera.htm (accessed 15 July 2006).
Dayton, P.K., Tegner, M.J., Edwards, P.B. & Riser, K.L. 1998. Sliding baselines, ghosts, and reduced expec-
tations in kelp forest communities. Ecological Applications 8, 309–322.
De Grave, S. & Whitaker, A. 1999. A census of maerl beds in Irish waters. Aquatic Conservation: Marine
and Freshwater Ecosystems 9, 303–311.
Delgado, O., Grau, A., Pou, S., Riera, F., Massuti, C., Zabala, M. & Ballesteros, E. 1997. Seagrass regression
caused by fish cultures in Fornells Bay (Menorca, western Mediterranean). Oceanologica Acta 20,
557–563.
396
Den Hartog, C. 1987. ‘Wasting disease’ and other dynamic phenomena in Zostera beds. Aquatic Botany 27,
3–14.
Desprez, M. 2000. Physical and biological impact of marine aggregate extraction along the French coast of
the Eastern English Channel: short- and long-term post-dredging restoration. ICES Journal of Marine
Science 57, 1428–1438.
de Villèle, X. & Verlaque, M. 1995. Changes and degradation in a Posidonia oceanica bed invaded by the
introduced tropical alga Caulerpa taxifolia in the northwestern Mediterranean. Botanica Marina 38,
79–87.
Diaz, R.J. 2001. Overview of hypoxia around the world. Journal of Environmental Quality 30, 275–280.
Dijkema, K.S. (ed.) 1984. Saltmarshes in Europe. Nature and Environment Series 30. Strasbourg, France:
Council of Europe.
Dodd, J. 2005. Native Oysters. Argyll, U.K.: Scottish Natural Heritage. Online. Available HTTP:
http://www.snh.org.uk (accessed 19 July 2006).
Donnan, D.W. & Moore, P.G. 2003. Introduction. Aquatic Conservation: Marine and Freshwater Ecosystems
13 (Suppl.), 1–3.
Doody, J.P. 2004. Coastal squeeze: a historical perspective. Journal of Coastal Conservation 10, 138 only.
Duarte, C.M. 2002. The future of seagrass meadows. Environmental Conservation 29, 192–206.
Dugan, P. (ed.) 1993. Wetlands in Danger. A World Conservation Atlas. New York: Oxford University Press.
EC. 2003. Interpretation Manual of European Union Habitats, Version EUR25. Brussels: EC, DG Environment.
Online. Available HTTP: http://ec.europa.eu/environment/nature/nature_conservation/eu_enlargement/
2004/pdf/habitats_im_en.pdf#search=‘interpretation%20manual%20Habitats%20directive’ (accessed
2 August 2006).
EC. 2004. Living with Coastal Erosion in Europe — Sediment and Space for Sustainability. Luxembourg:
OPOCE. Online. Available HTTP: http://www.eurosion.org/project/eurosion_en.pdf (accessed 11 Sep-
tember 2006).
Edgar, G.J., Barrett, N.S., Graddon, D.J. & Last, P.R. 2000. The conservation significance of estuaries: a
classification of Tasmanian estuaries using ecological, physical and demographic attributes as a case
study. Biological Conservation 92, 383–397.
EEA. 1998. Europe’s Environment: the Second Assessment. State of Environment report No 1/1998. Copen-
hagen: EEA. Online (summary). Available HTTP: http://reports.eea.europa.eu/92–828–3351–8/en
(accessed 24 June 2006).
EEA. 1999a. Coastal and marine zones. Chapter 3.14. Environment in the European Union at the Turn of the
Century. State of Environment report No 1/1999. Copenhagen: EEA. Online. Available HTTP:
http://reports.eea.eu.int/92–9157–202–0/en (accessed 4 August 2006).
EEA. 1999b. Corine Land Cover (CLC1990) 250 m - Version 06/1999. Online. Available HTTP: http://dataservice.
eea.europa.eu/atlas/viewdata/viewpub.asp?id=65 (accessed 4 August 2006).
EEA. 1999c. State and Pressures of the Marine and Coastal Mediterranean Environment. Environmental
Issues Series 5. Luxembourg: OPOCE. Online. Available HTTP: http://reports.eea.europa.eu/
ENVSERIES05/en/envissue05.pdf (accessed 4 August 2006).
EEA. 2001. Eutrophication in Europe’s Coastal Waters. Topic Report 7. Copenhagen: EEA. Online. Available
HTTP: http://reports.eea.eu.int/topic_report_2001_7 (accessed 4 August 2006).
EEA. 2002. Europe’s Biodiversity — Biogeographical Regions and Seas. Online. Available HTTP:
http://reports.eea.eu.int/report_2002_0524_154909/en (accessed 4 August 2006).
EEA. 2005. The European Environment — State and Outlook 2005. Copenhagen: EEA. Online. Available HTTP:
http://reports.eea.europa.eu/state_of_environment_report_2005_1/en/SOER2005_all.pdf (accessed
4 August 2006).
EEA. 2006a. The Changing Faces of Europe’s Coastal Areas. EEA Report 6/2006. Luxembourg: OPOCE.
Online. Available HTTP: http://reports.eea.europa.eu/eea_report_2006_6/en/eea_report_6_2006.pdf
(accessed 7 August 2006).
EEA. 2006b. Priority Issues in the Mediterranean Environment. EEA Report 4/2006. Luxembourg: OPOCE.
Online. Available HTTP: http://reports.eea.europa.eu/eea_report_2006_4/en/medsea_4_2006.pdf
397
Ekebom, J. & Erkkilä, A. 2003. Using aerial photography for identification of marine and coastal habitats
under the E.U.’s Habitats Directive. Aquatic Conservation — Marine and Freshwater Ecosystems 13,
287–304.
Eriksson, B.K. 2002. Long-Term Changes in Macroalgal Vegetation on the Swedish Coast. An Evaluation of
Eutrophication Effects with Special Emphasis on Increased Organic Sedimentation. Comprehensive
Summaries of Uppsala Dissertations from the Faculty of Science and Technology, Upsala University.
Uppsala, Sweden: Tryck & Medier.
Eriksson, B.K. & Bergstrom, L. 2005. Local distribution patterns of macroalgae in relation to environmental
variables in the northern Baltic Proper. Estuarine Coastal and Shelf Science 62, 109–117.
Eriksson, B.K., Johansson, G. & Snoeijs, P. 1998. Long-term changes in the sublitoral zonation of brown
algae in the southern Bothnian Sea. European Journal of Phycology 33, 241–249.
Eriksson, B.K., Johansson, G. & Snoeijs, P. 2002. Long-term changes in the macroalgal vegetation of the
inner Gullmar Fjord, Swedish Skagerrak coast. Journal of Phycology 38, 284–296.
EUCC — The Coastal Union. 1998. Posidonia beds. In Facts and Figures on Europe’s Biodiversity: State
and Trends 1998–1999, B. Delbaere (ed.). Technical Report Series. Tilburg: ECNC. Online. Available
HTTP: http://www.coastalguide.org/dune/posidi.html (accessed 7 August 2006).
Fanelli, G., Piraino, S., Belmonte, G., Geraci, S. & Boero, F. 1994. Human predation along Apulian rocky
coasts (SE Italy): desertification caused by Lithophaga lithophaga (Mollusca) fisheries. Marine Ecol-
ogy Progress Series 110, 1–8.
Finlayson, C.M. & Spiers, A.G. (eds) 1999. Global Review of Wetland Resources and Priorities for Wetland
Inventory. Supervising Scientist Report 144. Canberra: Supervising Scientist.
Franke, H.D. & Gutow, L. 2004. Long-term changes in the macrozoobenthos around the rocky island of
Helgoland (German Bight, North Sea). Helgoland Marine Research 58, 303–310.
Frid, C., Hammer, C., Law, R., Loeng, H., Pawlak, J.F., Reid, P.C. & Tasker, M. 2003. Environmental Status
of the European Seas. Copenhagen: ICES. Online. Available HTTP: http://www.bmu.de/files/pdfs/
allgemein/application/pdf/ices_report.pdf (accessed 7 August 2006).
Frid, C.L.J., Harwood, K.G., Hall, S.J. & Hall, J.A. 2000. Long-term changes in the benthic communities on
North Sea fishing grounds. ICES Journal of Marine Science 57, 1303–1309.
Glemaréc, M., LeFaou, Y. & Cuq, F. 1997. Long-term changes of seagrass beds in the Glenan Archipelago
(South Brittany). Oceanologica Acta 20, 217–227.
Gomoiu, M.-T. 1992. Marine eutrophication syndrome in the north-western part of the Black Sea. In Marine
Coastal Eutrophication, R.A. Vollenweider et al. (eds). Amsterdam: Elsevier, 683–692.
Grall, J. & Hall-Spencer, J.M. 2003. Problems facing maerl conservation in Brittany. Aquatic Conservation-
Marine and Freshwater Ecosystems 13(Suppl.), 55–64.
Gray, J.S. 1997. Marine biodiversity: patterns, threats and conservation needs. Biodiversity and Conservation
6, 153–175.
Green, E.P. & Short, F.T. 2003. World Atlas of Seagrasses. Berkeley, California: UNEP-WCMC, University
of California Press.
Green, M.J.B. & Paine, J. 1997. State of the world’s protected areas at the end of the twentieth century. Paper
presented at IUCN World Commission on Protected Areas Symposium on ‘Protected Areas in the
Twenty-First Century: From Islands to Networks’. Cambridge, U.K.: WCMC. Online. Available
HTTP: http://ariiprotejate.ngo.ro/docs/worldsap.pdf (accessed 28 July 2006).
Guidetti, P. 2001. Detecting environmental impacts on the Mediterranean seagrass Posidonia oceanica (L.)
Delile: the use of reconstructive methods in combination with ‘beyond BACI’ designs. Journal of
Guidetti, P., Fraschetti, S., Terlizzi, A. & Boero, F. 2003. Distribution patterns of sea urchins and barrens in
shallow Mediterranean rocky reefs impacted by the illegal fishery of the rock-boring mollusc Litho-
phaga lithophaga. Marine Biology 143, 1135–1142.
Günther, R.T. 1897. The oyster culture of the ancient Romans. Journal of the Marine Biological Association
of the United Kingdom 4, 360–365.
Hagen, N.T. 1995. Recurrent destructive grazing of successionally immature kelp forests by green sea-urchins
in Vestfjorden, northern Norway. Marine Ecology Progress Series 123, 95–106.
Hall-Spencer, J. 2005. Ban on maerl extraction. Marine Pollution Bulletin 50, 121 only.
398
Hall-Spencer, J.M., Grall, J., Moore, P.G. & Atkinson, R.J.A. 2003. Bivalve fishing and maerl-bed conservation
in France and the U.K. — retrospect and prospect. Aquatic Conservation: Marine and Freshwater
Ecosystems 13 (Suppl.), 33–41.
Hall-Spencer, J.M. & Moore, P.G. 2000. Scallop dredging has profound, long-term impacts on maerl habitats.
ICES Journal of Marine Science 57, 1407–1415.
Hamm, L., Capobianco, M., Dette, H.H., Lechuga, A., Spanhoff, R. & Stive, M.J.F. 2002. A summary of
European experience with shore nourishment. Coastal Engineering 47, 237–264.
Hannah, L., Lohse, D., Hutchinson, C., Carr, J.J. & Lankerani, A. 1994. A preliminary inventory of human
disturbance of world ecosystems. Ambio 23, 246–250.
Hauxwell, J., Cebrián, J., Furlong, C. & Valiela, I. 2001. Macroalgal canopies contribute to eelgrass (Zostera
marina) decline in temperate estuarine ecosystems. Ecology 82, 1007–1022.
Healy, M.G. & Hickey, K.R. 2002. Historic land reclamation in the intertidal wetlands of the Shannon estuary,
western Ireland. Journal of Coastal Research SI 36, 365–373.
Heck, K.L., Jr. & Crowder, L.B. 1991. Habitat structure and predator-prey interactions in vegetated aquatic
systems. In Habitat Structure: the Physical Arrangement of Objects in Space, S.S. Bell et al. (eds).
London: Chapman & Hall, 282–299.
HELCOM. 1998. Red List of Marine and Coastal Biotopes and Biotope Complexes of the Baltic Sea, Belt
Sea and Kattegat. Baltic Sea Environment Proceedings 75. Helsinki: HELCOM. Online. Available
HTTP: http://www.helcom.fi/stc/files/Publications/Proceedings/bsep75.pdf (accessed 7 August 2006).
Hemminga, M.A. & Duarte, C.M. 2000. Seagrass Ecology. Cambridge: Cambridge University Press.
Hiscock, K. & Kimmance, S. 2003. Review of Current and Historical Time-Series Seabed Biological Studies
in the U.K. and Near Europe. JNCC Report 336. Peterborough: JNCC. Online. Available HTTP:
http://www.jncc.gov.uk/pdf/jncc336.pdf (accessed 19 July 2006).
Hiscock, K., Sewell, J. & Oakley, J. 2005. Marine Health Check 2005. A Report to Gauge the Health of the
U.K.’s Sea-Life. Godalming: WWF-UK. Online. Available HTTP: http://www.wwf.org.uk/filelibrary/
pdf/marine_healthcheck05.pdf (accessed 7 August 2006).
Hiscock, K. & Tyler-Walters, H. 2006. Assessing the sensitivity of seabed species and biotopes — the Marine
Life Information Network (Marlin). Hydrobiologia 555, 309–320.
Hoffmann, R.C. 2005. A brief history of aquatic resource use in medieval Europe. Helgoland Marine Research
59, 22–30.
Holmer, M., Lassus, P., Stewart, J.E. & Wildish, D.J. 2001. ICES symposium on environmental effects of
mariculture. Introduction. ICES Journal of Marine Science 58, 363–368.
Hore, J.P. & Jex, E. 1880. The Deterioration of Oyster and Trawl Fisheries of England: Its Causes and Remedy.
London: Elliot Stock.
Hughes, R.G. & Paramor, O.A.L. 2004. On the loss of saltmarshes in south-east England and methods for
their restoration. Journal of Applied Ecology 41, 440–448.
ICES. 2006a. Report of the Working Group on Marine Habitat Mapping (WGMHM), 4–7 April, 2006, Galway,
Ireland. ICES CM 2006/MHC:05. Copenhagen: ICES. Online. Available HTTP: http://www.ices.dk/
reports/MHC/2006/WGMHM06.pdf (accessed 7 August 2006).
ICES. 2006b. Report of the Working Group on the Effects of Extraction of Marine Sediments on the Marine
Ecosystems (WGEXT), 4–7 April, 2006, Cork, Ireland. ICES CM 2006/MHC:07. Copenhagen: ICES.
Online. Available HTTP: http://www.ices.dk/reports/MHC/2006/WGMHM06.pdf (accessed 7 August
2006).
Islam, M.S. & Tanaka, M. 2004. Impacts of pollution on coastal and marine ecosystems including coastal and
marine fisheries and approach for management: a review and synthesis. Marine Pollution Bulletin 48,
624–649.
Jackson, A. 2003. Ostrea Edulis. Native Oyster. Marine Life Information Network: Biology and Sensitivity Key
Information Sub-programme. Plymouth, U.K.: Marine Biological Association of the United Kingdom.
Online. Available HTTP: http://www.marlin.ac.uk/species/Ostreaedulis.htm (accessed 7 August 2006).
Jackson, J.B.C., Kirby, M.X., Berger, W.H., Bjorndal, K.A., Botsford, L.W., Bourque, B.J., Bradbury, R.H.,
Cooke, R., Erlandson, J., Estes, J.A., Huges, T.P., Kidwell, S., Lange, C.B., Lenhian, H.S., Pandolfi,
J.M., Peterson, C.H., Steneck, R.S., Tegner, M.J. & Warner, R.R. 2001. Historical overfishing and the
recent collapse of coastal ecosystems. Science 293, 629–638.
399
Jansson, B.-O. & Dahlberg, K. 1999. The environmental status of the Baltic sea in the 1940s, today and in
the future. Ambio 28, 312–319.
Jaubert, J.M., Chisholm, J.R.M., Minghelli-Roman, A., Marchioretti, M., Morrow, J.H. & Ripley, H.T. 2003.
Re-evaluation of the extent of Caulerpa taxifolia development in the northern Mediterranean using
airborne spectrographic sensing. Marine Ecology Progress Series 263, 75–82.
Jeftic, L., Bernhard, M., Demetropulous, A., Fernex, F., Gabrielides, G.P., Gasparovic, F., Halim, Y., Orhon,
D. & Saliba, L.J. 1990. State of the Marine Environment in the Mediterranean Region. UNEP Regional
Seas Reports and Studies 132/1990 and MAP Technical Reports Series 28/1989. Athens: UNEP.
Online. Available HTTP: http://www.unepmap.org/home.asp (accessed 7 August 2006).
Jensen, A. 1998. The seaweed resources of Norway. In Seaweed Resources of the World, A.T. Critchley & M.
Ohno (eds). Yokosuka, Japan: Japan International Cooperation Agency, 200–209.
Johansson, G., Eriksson, B.K., Pedersén, M. & Snoeijs, P. 1998. Long-term changes of macroalgal vegetation
in the Skagerrak area. Hydrobiologia 385, 121–138.
Johnson, K.A. 2002. A Review of National and International Literature on the Effects of Fishing on Benthic
Habitats. NOAA Technical memorandum. NMFS-F/SPO-57. Silver Spring, Maryland: U.S. Depart-
ment of Commerce, National Marine Fisheries Service, NOAA. Online. Available HTTP: http://www.
nmfs.noaa.gov/habitat/habitatprotection/pdf/efh/literature/KJohnson.pdf (accessed 7 August 2006).
Jones, T.A. & Hughes, J.M.R. 1993. Wetland inventories and wetland loss studies: a European perspective.
In Waterfowl and Wetland Conservation in the 1990s: A Global Perspective, M. Moser et al. (eds).
Proceedings of the IWRB Symposium, St. Petersburg, Florida. November 1992. IWRB Special
Publication No. 26. Slimbridge, U.K.: IWRB, 164–169.
Kabat, P.W., van Vierssen, J., Veraart, J., Vellinga, P. & Aerts, J. 2005. Climate proofing the Netherlands.
Nature 438, 283–284.
Kangas, P., Autio, H., Hällfors, G., Luther, H., Niemi, Å. & Salemaa, H. 1982. A general model of the decline of
Fucus vesiculosus at Tvärminne, south coast of Finland in 1977–1981. Acta Botanica Fennica 118, 1–27.
Karlson, K., Rosenberg, R. & Bonsdorff, E. 2002. Temporal and spatial large-scale effects of eutrophication
and oxygen deficiency on benthic fauna in Scandinavian and Baltic waters — a review. Oceanography
and Marine Biology: An Annual Review 40, 427–489.
Kautsky, N., Kautsky, H., Kautsky, U. & Wærn, M. 1986. Decreased depth penetration of Fucus vesiculosus
(L.) since the 1940s indicates eutrophication of the Baltic Sea. Marine Ecology Progress Series 28, 1–8.
Keddy, P.A. 2000. Wetland Ecology. Principles and Conservation. Cambridge: Cambridge University Press.
Kelleher, G., Bleakley, C. & Wells, S. (eds) 1995. A Global Representative System of Marine Protected Areas.
Volume 1. Antarctic, Arctic, Mediterranean, Northwest Atlantic, Northeast Atlantic and Baltic. Wash-
ington, D.C.: The Great Barrier Reef Marine Park Authority, The World Bank, The World Conservation
Union. Online. Available HTTP: http://www.deh.gov.au/coasts/mpa/nrsmpa/global/volume1/#download
(accessed 28 July 2006).
Kennedy, R.J. & Roberts, D. 1999. A survey of the current status of the flat oyster Ostrea edulis in Strangford
Lough, Northern Ireland, with a view to the restoration of its oyster beds. Biology and Environment:
Proceedings of the Royal Irish Academy 99B, 79–88.
Kennish, M.J. 2002. Environmental threats and environmental future of estuaries. Environmental Conservation
29, 78–107.
Knottnerus, O.S. 2005. History of human settlement, cultural change and interference with the marine
environment. Helgoland Marine Research 59, 2–8.
Korringa, P. 1952. Recent advances in oyster biology. The Quarterly Review of Biology 27, 266–308.
Laffoley, D.d. 2000. Historical Perspective and Selective Review of the Literature on Human Impacts on the
U.K.’s Marine Environment. English Nature Research Reports No. 391. Peterbrorough, U.K.: English
Nature for the DETR Working Group on the Review of Marine Nature Conservation. Online. Available
HTTP: http://www.jncc.gov.uk/pdf/ENRR391.pdf (accessed 4 August 2006).
Lee, M. 2001. Coastal defence and the Habitats Directive: predictions of habitat change in England and Wales.
The Geographical Journal 167, 39–56.
Leslie, H., Ruckelshaus, M., Ball, I., Andelman, S. & Possingham, H. 2003. Using siting algorithms in the
design of marine reserve networks. Ecological Applications 13, 185–198.
400
Lindeboom, H.J. 1995. Protected areas in the North Sea: an absolute need for future marine research. Helgoland
Marine Research 49, 591–602.
Lindeboom, H.J. & de Groot, S.J. (eds) 1998. IMPACT-II. The Effects of Different Types of Fisheries on the
North Sea and Irish Sea Benthic Ecosystems. Texel, The Netherlands: Netherlands Institute for Sea
Research.
Lotze, H.K. 2004. Repetitive history of resource depletion and mismanagement: the need for a shift in
perspective. Marine Ecology Progress Series 274, 269–303.
Lotze, H.K. 2005. Radical changes in the Wadden Sea fauna and flora over the last 2000 years. Helgoland
Marine Research 59, 71–83.
Lotze, H.K., Lenihan, H.S., Bourque, B.J., Bradbury, R.H., Cooke, R.G., Kay, M.C., Kidwell, S.M., Kirby,
M.X., Peterson, C.H. & Jackson, J.B.C. 2006. Depletion, degradation, and recovery potential of
estuaries and coastal seas. Science 312, 1806–1809.
Lotze, H.K. & Reise, K. 2005. Ecological history of the Wadden Sea. Helgoland Marine Research 59, 1 only.
Lotze, H.K., Reise, K., Worm, B., Van Beusekom, J., Busch, M., Ehlers, A., Heinrich, D., Hoffmann, R.C.,
Holm, P., Jensen, C., Knottnerus, O.S., Langhanki, N., Prummel, W., Vollmer, M. & Wolff, W.J. 2005.
Human transformations of the Wadden Sea ecosystem through time: a synthesis. Helgoland Marine
Research 59, 84–95.
Lozan, J.L. 1996. Nutzung und Veraenderungen von Mollusken und Krebsen der Ostsee. In Warnsignale aus
der Ostsee, J.L. Lozan et al. (eds). Berlin: Parey, 197–201.
Lumb, C.M. 1990. Algal depth distributions and long-term turbidity changes in the Menai Strait, North Wales.
Progress in Underwater Science 15, 85–99.
Lundälv, T., Larsson, C.S. & Axelsson, L. 1986. Long-term trends in algal-dominated rocky subtidal commu-
nities on the Swedish west coast — a transitional system? Hydrobiologia 142, 81–95.
Lüning, K. 1990. Seaweeds, Their Environment, Biogeography and Ecophysiology. New York: John Wiley &
Sons.
Mackenzie, C.L., Jr., Burrell, V.G., Jr., Rosenfield, A. & Hobart, W.L. (eds). 1997. The History, Present
Condition and Future of the Molluscan Fisheries of North and Central America and Europe. Volume 3,
Europe. NOAA Technical Report NMFS, 129. Seattle, Washington: U.S. Department of Commerce.
Marbà, N., Duarte, C.M., Cebrian, J., Enríquez, S., Gallegos, M.E., Olesen, B. & Sand-Jensen, K. 1996.
Growth and population dynamics of Posidonia oceanica on the Spanish Mediterranean coast: eluci-
dating seagrass decline. Marine Ecology Progress Series 137, 203–213.
Marbà, N., Duarte, C.M., Holmer, M., Martínez, R., Basterretxea, G., Orfila, A., Jordi, A. & Tintore, J. 2002.
Effectiveness of protection of seagrass (Posidonia oceanica) populations in Cabrera National Park.
Environmental Conservation 29, 509–518.
Marchetti, R., Provini, A. & Crosa, G. 1989. Nutrient load carried by the river Po into the Adriatic Sea. Marine
Pollution Bulletin 20, 168–172.
Martin, D., Bertasi, F., Colangelo, M.A., de Vries, M., Frost, M., Hawkins, S.J., Macpherson, E., Moschella,
P.S., Satta, M.P., Thompson, R.C. & Ceccherelli, V.U. 2005. Ecological impact of coastal defence
structures on sediments and mobile infauna: evaluating and forecasting consequences of unavoidable
modifications of native habitats. Coastal Engineering 52, 1027–1051.
McEvedy, C. & Jones, R. 1978. Atlas of World Population History. New York: Penguin.
Meinesz, A., Belsher, T., Thibaut, T., Antolic, B., Mustapha, K.B., Boudouresque, C.-F., Chiaverini, D., Cinelli,
F., Cottalorda, J.-M., Djellouli, A., El Abed, A., Orestano, C., Grau, A.M., Ivesa, L., Jaklin, A., Langar,
H., Massuti-Pascual, E., Peirano, A., Tunesi, L., de Vaugelas, J., Zavodnik, N. & Zuljevic, A. 2001.
The introduced green alga Caulerpa taxifolia continues to spread in the Mediterranean. Biological
Invasions 3, 201–210.
Meinesz, A., Lefevre, J.R. & Astier, J.M. 1991. Impact of coastal development on the infralittoral zone along
the southeastern Mediterranean shore of continental France. Marine Pollution Bulletin 23, 343–347.
Messner, U. & von Oertzen, J.A. 1991. Long-term changes in the vertical distribution of macrophytobenthic
communities in the Greifswalder Bodden. Acta Ichthyologica et Piscatoria 21(Suppl.), 135–143.
Middelboe, A.L. & Sand-Jensen, K. 2000. Long-term changes in macroalgal communities in a Danish estuary.
401
Milazzo, M., Badalamenti, F., Ceccherelli, G. & Chemello, R. 2004. Boat anchoring on Posidonia oceanica
beds in a Marine Protected Area (Italy, western Mediterranean): effect of anchor types in different
anchoring stages. Journal of Experimental Marine Biology and Ecology 299, 51–62.
Minello, T., Zimmerman, R. & Medina, R. 1994. The importance of edge for natant macrofauna in a created
salt marsh. Wetlands 14, 184–198.
Mock, G., White, R. & Wagener, A. 1998. Farming Fish: the Aquaculture Boom, W. Vanasselt (ed.). Originally
written for World Resources 1998–1999, updated for EarthTrends (July 2001). Online. Available
HTTP: http://earthtrends.wri.org/features/view_feature.cfm?theme=1&fid=20 (accessed 20 July 2006).
Morris, J.T., Sundareshwar, P.V., Nietch, C.T. & Kjerfve, B.C.D.R. 2002. Responses of coastal wetlands to
rising sea level. Ecology 83, 2869–2877.
Moschella, P.S., Abbiati, M., Åberg, P., Airoldi, L., Anderson, J.M., Bacchiocchi, F., Bulleri, F., Dinesen, G.E.,
Frost, M., Gacia, E., Granhag, L., Jonsson, P.R., Satta, M.P., Sundelof, A., Thompson, R.C. & Hawkins,
S.J. 2005. Low-crested coastal defence structures as artificial habitats for marine life: using ecological
criteria in design. Coastal Engineering 52, 1053–1071.
MPA Global. 2006. A database of the world’s Marine Protected Areas. University of British Columbia. Online.
Available HTTP: http//www.mpaglobal.org (accessed 9 August 2006).
Munda, I.M. 1993. Changes and degradation of seaweed stands in the northern Adriatic. Hydrobiologia
260/261, 239–253.
Munda, I.M. 2000. Long-term marine floristic changes around Rovinj (Istrian coast, north Adriatic) estimated
on the basis of historical data from Paul Kuckuck’s field diaries from the end of the nineteenth century.
Nehring, D. 1992. Eutrophication in the Baltic Sea. In Marine Coastal Eutrophication, R.A. Vollenweider
et al. (eds). Amsterdam: Elsevier, 673–682.
Newell, R.C., Seiderer, L.J. & Hitchcock, D.R. 1998. The impact of dredging works in coastal waters: a review
of the sensitivity to disturbance and subsequent recovery of biological resources on the seabed.
Nicholls, R.J., Hoozemans, F.M.J. & Marchand, M. 1999. Increasing flood risk and wetland losses due to
global sea-level rise: regional and global analyses. Global Environmental Change — Human and
Policy Dimensions 9 (Suppl.), 69–87.
Nilsson, J., Engkvist, R. & Persson, L.E. 2004. Long-term decline and recent recovery of Fucus populations
along the rocky shores of southeast Sweden, Baltic Sea. Aquatic Ecology 38, 587–598.
Nivet, C. & Frazier, S. 2004. A Review of European Wetland Inventory Information. Almere The Netherlands:
RIZA, Evers Litho en Druk.
Occhipinti-Ambrogi, A. 2001. Transfer of marine organisms: a challenge to the conservation of coastal
biocenoses. Aquatic Conservation: Marine and Freshwater Ecosystems 11, 243–251.
Ocean Studies Board. 2004. Nonnative Oysters in the Chesapeake Bay. Washington, D.C.: National Academy
Press.
Olenin, S. & Klovaité, K. 1998. Introduction to the marine and coastal environments of Lithuania. In Red List
of Marine and Coastal Biotopes and Biotope Complexes of the Baltic Sea, Belt Sea and Kattegat. Baltic
Sea Environment Proceedings 75. Helsinki: HELCOM. Online. Available HTTP: http://www.helcom.fi/
stc/files/Publications/Proceedings/bsep75.pdf (accessed 7 August 2006).
OSPAR Commission. 2005. Case Reports for the Initial List of Threatened and/or Declining Species and Habitats
in the OSPAR Maritime Area. Biodiversity Series: OSPAR Commission. Online. Available HTTP:
http://www.ospar.org/documents/dbase/publications/p00198_Case%20reports%20for%20Initial%20list
%20of%20species%20and%20habitats%202005%20version.pdf (accessed 8 August 2006).
Pandolfi, J.M., Bradbury, R.H., Sala, E., Hughes, T.P., Bjorndal, K.A., Cooke, R.G., McArdle, D., McClena-
chan, L., Newman, M.J.H., Paredes, G., Warner, R.R. & Jackson, J.B.C. 2003. Global trajectories of
the long-term decline of coral reef ecosystems. Science 301, 955–958.
Pasqualini, V., Pergent-Martini, C., Clabaut, P. & Pergent, G. 1998. Mapping of Posidonia oceanica using
aerial photographs and side scan sonar: application off the island of Corsica (France). Estuarine
Coastal and Shelf Science 47, 359–367.
Pedersén, M. & Snoeijs, P. 2001. Patterns of macroalgal diversity, community composition and long-term
changes along the Swedish west coast. Hydrobiologia 459, 83–102.
402
Petersen, K.S., Rasmussen, K.L., Heinemeier, J. & Rud, N. 1992. Clams before Columbus? Nature 359, 679
only.
Piazzi, L., Acunto, S. & Cinelli, F. 2000a. Mapping of Posidonia oceanica beds around Elba Island (western
Mediterranean) with integration of direct and indirect methods. Oceanologica Acta 23, 339–346.
Piazzi, L., Acunto, S., Papi, I., Pardi, G. & Cinelli, F. 2000b. Mapping of the seagrasses beds in Tuscany
(Italy): situation in 1998. Biologia Marina Mediterranea 7, 594–596.
Pihl, L., Baden, S., Kautsky, N., Ronnback, P., Soderqvist, T., Troell, M. & Wennhage, H. 2006. Shift in fish
assemblage structure due to loss of seagrass Zostera marina habitats in Sweden. Estuarine Coastal
and Shelf Science 67, 123–132.
Possingham, H., Ball, I. & Andelman, S. 2002. Mathematical models for identifying representative reserve
networks. In Quantitative Methods in Conservation Biology, S. Ferson & M.A. Burgman (eds). New
York: Springer-Verlag, second edition, 291–306.
Pruett, L. & Cimino, J. 2000. Global Maritime Boundaries Database (GMBD). Fairfax, Virginia, U.S.:
Veridian — MRJ Technology Solutions. Online. Available HTTP: http://earthtrends.wri.org/text/
coastal-marine/variable-61.html (accessed 13 July 2006).
Pukaric, S.B.G.W. & Jorissen, F.J. 1990. Successive appearance of subfossil phytoplankton species in Holocene
sediments of the northern Adriatic and its relation to the increased eutrophication pressure. Estuarine
Coastal and Shelf Science 31, 177–187.
Rask, N., Pedersen, S.E. & Jensen, M.H. 1999. Response to lowered nutrient discharge in coastal waters
around the island of Funen, Denmark. Hydrobiologia 393, 69–81.
Reise, K. 1994. Changing life under the tides of the Wadden Sea during the twentieth century. Ophelia Suppl.
6, 117–125.
Reise, K. 2005. Coast of change: habitat loss and transformations in the Wadden Sea. Helgoland Marine
Research 59, 9–21.
Reise, K., Olenin, S. & Thieltges, D.W. 2006. Are aliens threatening European aquatic coastal ecosystems?
Rippon, S. 1997. The Severn Estuary: Landscape Evolution and Wetland Reclamation. London: Leicester
University Press.
Rippon, S. 2000. The Transformation of Coastal Wetlands: Exploitation and Management of Marshland
Landscapes in North West Europe During the Roman and Medieval Periods. London: British Academy.
Rismondo, A., Guidetti, P. & Curiel, D. 1997. Presenza delle fanerogame marine nel Golfo di Venezia: un
aggiornamento. Bollettino del Museo Civico di Storia Naturale di Venezia 47, 317–328.
Rodríguez-Prieto, C. & Polo, L. 1996. Effects of sewage pollution in the structure and dynamics of the
community of Cystoseira mediterranea (Fucales, Phaeophyceae). Scientia Marina 60, 253–263.
Rosenberg, R., Elmgren, R., Fleischer, S., Jonsson, P., Persson, G. & Dahlin, H. 1990. Marine eutrophication
case studies in Sweden. Ambio 19, 102–108.
Russo, G.F. & Cicogna, F. 1991. The date mussel (Lithophaga lithophaga), a ‘case’ in the Gulf of Naples. In
Les Espèces Marines à Protéger en Méditérranée, C.F. Boudouresque et al. (eds). Marseille: GIS
Posidonie, 141–150.
Sala, E. 2004. The past and present topology and structure of Mediterranean subtidal rocky-shore food webs.
Ecosystems 7, 333–340.
Sandulli, R., Bianchi, C.N., Cocito, S., Morgigni, M., Sgorbini, S., Silvestri, C., Morri, C. & Peirano, A. 1994.
Status of some Posidonia oceanica meadows on the Ligurian coast influenced by the Haven oil spill.
In Atti Del 10° Congresso AIOL (Associazione Italiana Oceanologia e Linmologia), G. Albertelli
et al. (eds). Genova, Italy: AIOL, 277–286.
Sangiorgi, F. & Donders, T.H. 2004. Reconstructing 150 years of eutrophication in the north-western Adriatic
Sea (Italy) using dinoflagellate cysts, pollen and spores. Estuarine Coastal and Shelf Science 60,
69–79.
Schories, D., Albrecht, A. & Lotze, H. 1997. Historical changes and inventory of macroalgae from Königshafen
Bay in the northern Wadden Sea. Helgoländer Meeresuntersuntersuchungen 51, 321–341.
Sebens, K.P. 1994. Biodiversity of coral reefs: what are we loosing and why? American Zoologist 34, 115–133.
Sfriso, A. 1987. Flora and vertical distribution of macroalgae in the lagoon of Venice: a comparison with
previous studies. Giornale Botanico Italiano 121, 69–85.
403
Sfriso, A. & La Rocca, B. 2005. Aggiornamento sulle macroalghe presenti lungo i litorali e sui bassofondali
della laguna di Venezia. Lavori della Società Veneziana di Scienze Naturali 30, 45–56.
Short, F.T. & Wyllie-Echeverria, S. 1996. Natural and human-induced disturbance of seagrasses. Environmental
Conservation 23, 17–27.
Sih, A., Jonsson, B.G. & Luikart, G. 2000. Habitat loss: ecological, evolutionary and genetic consequences.
Simonini, R., Ansaloni, I., Bonvicini Pagliai, A.M., Cavallini, F., Iotti, M., Mauri, M., Montanari, G., Preti,
M., Rinaldi, A. & Prevedelli, D. 2005. The effects of sand extraction on the macrobenthos of a relict
sands area (northern Adriatic Sea): results 12 months post-extraction. Marine Pollution Bulletin 50,
768–777.
Spalding, M.D., Green, E.P. & Rvilious, C. 2001. World Atlas of Coral Reefs. Berkeley, California: UNEP-
WCMC, University of California Press.
Stanners, D. & Bourdeau, P. (eds) 1995. Europe’s Environment. The DOBRIS Assessment. State of Environment
Report 1/1995. Copenhagen: EEA. Online. Available HTTP: http://reports.eea.eu.int/92–826–5409–5/en
Steneck, R.S., Graham, M.H., Bourque, B.J., Corbett, D., Erlandson, J.M., Estes, J.A. & Tegner, M.J. 2002.
Kelp forest ecosystems: biodiversity, stability, resilience and future. Environmental Conservation 29,
436–459.
Suchanek, T.H. 1994. Temperate coastal marine communities: biodiversity and threats. American Zoologist
34, 100–114.
Thibaut, T., Pinedo, S., Torras, X. & Ballesteros, E. 2005. Long-term decline of the populations of Fucales
(Cystoseira spp. and Sargassum spp.) in the Alberes coast (France, north-western Mediterranean).
Marine Pollution Bulletin 50, 1472–1489.
Thompson, R.C., Crowe, T.P. & Hawkins, S.J. 2002. Rocky intertidal communities: past environmental
changes, present status and predictions for the next 25 years. Environmental Conservation 29, 168–191.
Thrush, S.F. & Dayton, P.K. 2002. Disturbance to marine benthic habitats by trawling and dredging: impli-
cations for marine biodiversity. Annual Reviews of Ecology and Systematics 33, 449–473.
Trombini, C., Fabbri, D., Lombardo, M., Vassura, I., Zavoli, E. & Horvat, M. 2003. Mercury and methylm-
ercury contamination in surficial sediments and clams of a coastal lagoon (Pialassa Baiona, Ravenna,
Italy). Continental Shelf Research 23, 1821–1831.
Tudela, S. 2004. Ecosystem Effects of Fishing in the Mediterranean: An Analysis of the Major Threats of
Fishing Gear and Practices to Biodiversity and Marine Habitats. Studies and Reviews. General
Fisheries Commission for the Mediterranean 74. Rome: FAO. Online. Available HTTP: http://www.
fao.org/docrep/007/y5594e/y5594e00.htm (accessed 1 August 2006).
Turner, S.J., Thrush, S.F., Hewitt, J.E., Cummings, V.J. & Funnell, G. 1999. Fishing impacts and the degradation
or loss of habitat structure. Fisheries Management and Ecology 6, 401–420.
Tyler-Walters, H. 2001. Ostrea Edulis Beds on Shallow Sublittoral Muddy Sediment. Marine Life Information
Network: Biology and Sensitivity Key Information Sub-programme. Plymouth, U.K.: Marine Biolog-
ical Association of the United Kingdom. Online. Available HTTP: http://www.marlin.ac.uk/biotopes/
Bio_BasicInfo_IMX.Ost.htm (accessed 8 August 2006).
U.K. Biodiversity Group. 1999. U.K. Biodiversity Group Tranche 2 Action Plans — Volume 5: Maritime
Species and Habitats. Peterborough, U.K.: English Nature. Online. Available HTTP: http://www.
ukbap.org.uk/Library/Tranche2_Vol5.pdf (accessed 2 August 2006).
U.K. Steering Group. 1995. Biodiversity: The U.K. Steering Group Report — Volume 2: Action Plans (Annex F:
Lists of Key Species, Key Habitats and Broad Habitats). London: HMSO. Online. Available HTTP:
http://www.ukbap.org.uk/Library/Tranche1_Ann_f.pdf (accessed 2 August 2006).
UNEP/FAO/WHO. 1996. Assessment of the State of Eutrophication in the Mediterranean Sea. MAP Technical
Series 106. Athens: UNEP.
UNEP/MAP/PAP. 2001. White Paper: Coastal Zone Management in the Mediterranean. Split: Priority
Actions Programme. Online. Available HTTP: http://www.pap-thecoastcentre.org/pdfs/ICAM%20in
%20Mediterranean%20%20White%20Paper.pdf#search=‘White%20paper%3A%20coastal%20zon%
20management%20in%20the%20Mediterranean’ (accessed 8 August 2006).
404
UNEP/WCMC. 2006. 2006 World Database on Protected Areas: UNEP-WCMC. Online. Available HTTP:
http://sea.unep-wcmc.org/wdpa/ (accessed 8 August 2006).
Valiela, I. 2006. Global Coastal Change. Singapore: Blackwell.
Valiela, I., Bowen, J.L. & York, J.K. 2001. Mangrove forests: one of the world’s threatened major tropical
environments. BioScience 51, 807–815.
van Beusekom, J.E.E. 2005. A historic perspective on Wadden Sea eutrophication. Helgoland Marine Research
59, 45–54.
van Dalfsen, J.A., Essink, K., Madsen, H.T., Birklund, J., Romero, J. & Manzanera, M. 2000. Differential
response of macrozoobenthos to marine sand extraction in the North Sea and the western Mediterra-
nean. ICES Journal of Marine Science, 57, 1439–1445.
van den Berg, J.B., Kozyreff, G., Lin, H.-X., McDarby, J., Peletier, M.A., Planqué, R. & Wilson, P.L. 2005.
Japanese oysters in Dutch waters. Nieuw Archief voor Wiskunde 5/6, 130–141.
Vogt, H. & Schramm, W. 1991. Conspicuous decline of Fucus in Kiel Bay (western Baltic): what are the
causes? Marine Ecology Progress Series 69, 189–194.
Váradi, L. 2001. Review of trends in the development of European inland aquaculture linkages with fisheries.
Fisheries Management and Ecology 8, 453–462.
Watling, L. & Norse, E.A. 1998. Disturbance of the seabed by mobile fishing gear: a comparison to forest
clearcutting. Conservation Biology 12, 1180–1197.
Wilcove, D.S., Rothstein, D., Dobow, J., Phillips, A. & Losos, E. 1998. Quantifying threats to imperiled
species in the United States. BioScience 48, 607–615.
Wilkie, M.L. & Fortuna, S. 2003. Status and Trends in Mangrove Area Extent Worldwide. Forest Resources
Assessment Working Paper 63. Rome: Forest Resources Division, FAO. Online. Available HTTP:
http://www.fao.org/docrep/007/j1533e/j1533e00.HTM (accessed 4 August 2006).
Wilkinson, C.R. (ed.) 2004. Status of Coral Reefs of the World: 2004. Volumes 1 and 2. Townsville, Australia:
Australian Institute of Marine Science. Online. Available HTTP: http://www.aims.gov.au/pages/
research/coral-bleaching/scr2004/index.html (accessed 4 August 2006).
Wilson, S., Blake, C., Berges, J.A. & Maggs, C.A. 2004. Environmental tolerances of free-living coralline
algae (maerl): implications for European marine conservation. Biological Conservation, 120, 283–293.
Wolff, W.J. 1992. The end of a tradition: 1000 years of embankment and reclamation of wetlands in the
Netherlands. Ambio 21, 287–291.
Wolff, W.J. 1997. Development of the conservation of Dutch coastal waters. Aquatic Conservation: Marine
and Freshwater Ecosystems 7, 165–177.
Wolff, W.J. 2000. Causes of extirpations in the Wadden Sea, an estuarine area in the Netherlands. Conservation
Biology 14, 876–885.
Wolff, W.J. 2005. The exploitation of living resources in the Dutch Wadden Sea: a historical overview.
Wolters, M., Bakker, J.P., Bertness, M.D., Jefferies, R.L. & Moller, I. 2005. Saltmarsh erosion and restoration
in south-east England: squeezing the evidence requires realignment. Journal of Applied Ecology 42,
844–851.
Worm, B., Lotze, H.K., Boström, C., Engkvist, R., Labanauskas, V. & Sommer, U. 1999. Marine diversity
shift linked to interactions among grazers, nutrients and propagule banks. Marine Ecology Progress
Series 185, 309–314.
Yonge, C.M. 1966. Oysters. London: Collins.
Zaitsev, Y.P. (ed.) 2006. Black Sea Red Data Book. Online. Available HTTP: http://www.grid.unep.ch/bsein/
redbook/index.htm (accessed 11 July 2006).
Zavodnik, N. & Jaklin, A. 1990. Long-term changes in the northern Adriatic marine phanerogam beds. Rapport
de la Commission Internationale pour l’Exploration Scientifique de la Mer Méditerranée 32, 15 only.
405

Taylor & Francis
CLIMATE CHANGE AND AUSTRALIAN MARINE LIFE

E.S. POLOCZANSKA1, R.C. BABCOCK2, A. BUTLER1, A.J. HOBDAY3,6,
O. HOEGH-GULDBERG4, T.J. KUNZ3, R. MATEAR3, D.A. MILTON1,
T.A. OKEY1 & A.J. RICHARDSON1,5
1Wealth from Oceans Flagship — CSIRO Marine & Atmospheric Research, PO Box 120, Cleveland,
Queensland 4163, Australia

E-mail: elvira.poloczanska@csiro.au
2Wealth from Oceans Flagship — CSIRO Marine & Atmospheric Research, Private Bag 5,
Floreat, Western Australia 6913, Australia

3Wealth from Oceans Flagship — CSIRO Marine & Atmospheric Research, GPO Box 1538,
Hobart, Tasmania 7001, Australia

4University of Queensland, Centre for Marine Studies,
St Lucia, Queensland 4072, Australia

5University of Queensland, Department of Mathematics,
St Lucia, Queensland 4072, Australia

6University of Tasmania, School of Zoology, Private Bag 5,
Hobart, Tasmania 7001, Australia
Abstract Australia’s marine life is highly diverse and endemic. Here we describe projections of
climate change in Australian waters and examine from the literature likely impacts of these changes
on Australian marine biodiversity. For the Australian region, climate model simulations project oceanic
warming, an increase in ocean stratification and decrease in mixing depth, a strengthening of the
East Australian Current, increased ocean acidification, a rise in sea level, alterations in cloud cover
and ozone levels altering the levels of solar radiation reaching the ocean surface, and altered storm
and rainfall regimes. Evidence of climate change impacts on biological systems are generally scarce
in Australia compared to the Northern Hemisphere. The poor observational records in Australia are
attributed to a lack of studies of climate impacts on natural systems and species at regional or
national scales. However, there are notable exceptions such as widespread bleaching of corals on
the Great Barrier Reef and poleward shifts in temperate fish populations. Biological changes are
likely to be considerable and to have economic and broad ecological consequences, especially in
climate-change ‘hot spots’ such as the Tasman Sea and the Great Barrier Reef.
Introduction
The global climate is changing and is projected to continue changing at a rapid rate for the next
100 yr (IPCC 2001, 2007). Average global temperatures have risen by 0.6 ± 0.2°C over the twentieth
century and this warming is likely to have been greater than for any other century in the last
millennium. The 1990s were the warmest decade globally of the past century; and the present
decade may be warmest yet (Hansen et al. 2006). Most of the warming observed during the last
50 yr is attributable to anthropogenic forcing by greenhouse gas emissions (Karoly & Stott 2006).
The increase in global temperature is likely to be accompanied by alterations in patterns and strength
of winds and ocean currents, atmospheric and ocean stratification, a rise in sea levels, acidification
of the oceans and changes in rainfall, storm patterns and intensity. Evidence is mounting that the
407
E.S. POLOCZANSKA ET AL.
changing climate is already impacting terrestrial, marine and freshwater ecosystems (Hoegh-
Guldberg 1999, Walther et al. 2002, Parmesan & Yohe 2003, Root et al. 2003, Walther et al. 2005).
Species’ distributions are shifting poleward (Parmesan et al. 1999, Thomas & Lennon 1999,
Beaugrand et al. 2002, Hickling et al. 2006), plants are flowering earlier and growing seasons are
lengthening (Edwards & Richardson 2004, Wolfe et al. 2005, Linderholm 2006, Schwartz et al.
2006) and timing of peak breeding and migrations of animals are altering (Both et al. 2004,
Lehikoinen et al. 2004, Weishampel et al. 2004, Jonzén et al. 2006, Menzel et al. 2006). Most of
this evidence, however, is from the Northern Hemisphere, with few examples from the Southern
Hemisphere and only a handful from Australia (Chambers 2006). The lack of observations in
Australia is attributed to a lack of studies of climate impacts on natural systems and species at
regional or national scales. Further, the extent of historical biological datasets in Australia is largely
unknown, many are held by small organisations or by individuals and the value of these datasets
may not be recognised (Chambers 2006).
Because of the unique geological, oceanographic and biological characteristics of Australia,
conclusions from climate impact studies in the Northern Hemisphere are not easily transferable to
Australian systems. Including fringing islands, Australia has a coastline of almost 60,000 km
(Figure 1) that spans from southern temperate waters of Tasmania and Victoria (~45°S) to northern
tropical waters of Cape York, Queensland (~10°S). Australia is truly a maritime country with over
90% of the population living within 120 km of the coast. Most of Australia’s population of 20 million
live in the southeast with the west and north coasts being sparsely populated. Around 40% of
Australia’s population live in the cities of Sydney and Melbourne alone (Australian Bureau of
Statistics 2006).
10°
Exmouth Torres Strait
Gulf Darwin Gulf of
Indian Scott
Carpentaria
Ocean Reef Cape
York Great Barrier Reef
20°
Hervey Bay Pacific Ocean
Australia
Shark Bay
Brisbane Moreton Bay
Houtman Abrolhos
30° Islands
Hawkesbury Estuary
Perth Adelaide Sydney Botany Bay
Albany
Melbourne
Corner Inlet New Zealand
Bass Strait Tasman Sea
40° Tasmania
Hobart
Southern Ocean Tasmanian Seamounts

Marine Reserve
50°
110° 120° 130° 140° 150° 160° 170° 180° 190°
Figure 1 (See also Colour Figure 1 in the insert following page 344.) Map of Australia indicating the locations
discussed in the text. The 200 nm EEZ for Australia is marked by the dashed line, and the 200 m depth contour
by the solid line.
408
Australia has sovereign rights over ~8.1 million km2 of ocean and this area generates consid-
erable economic wealth estimated as $A52 billion per year or about 8% of gross domestic product
(CSIRO 2006). Fisheries and aquaculture are important industries in Australia, both economically
(gross value over $A2.5 billion) and socially. Marine life and ecosystems also provide invaluable
services including coastal defence, nutrient recycling and greenhouse gas regulation valued globally
at $US 22 trillion ($A27 trillion) per annum (Costanza et al. 1997). The annual economic values
of Australian marine biomes have been estimated: open ocean $A464.7 billion, seagrass/algal beds
$A175.1 billion, coral reefs $A53.5 billion, shelf system $A597.9 billion and tidal marsh/mangroves
$A39.1 billion (Blackwell 2005). This assessment assumes Australian marine ecosystems are
unstressed so actual values may be lower for degraded systems. Compared to other countries,
relatively little is known about the biology and ecology of Australia’s maritime realm, mainly due
to the inaccessibility and remoteness of much of the coast as highlighted by the discovery of living
stromatolites (representing the one of the oldest known forms of life on Earth) in Western Australia
in the 1950s (Logan 1961).
Australia is unique among continents in that both the west and east coasts are bounded by
major poleward-flowing warm currents (Figure 2), which have considerable influence on marine
flora and fauna. The East Australian Current (EAC) originates in the Coral Sea and flows southward
before separating from the continental margin to flow northeast and eastward into the Tasman Sea
(Ridgway & Godfrey 1997, Ridgway & Dunn 2003). Eddies spawned by the EAC continue
southward into the Tasman Sea bringing episodic incursions of warm water to temperate eastern
Australia and Tasmanian waters (Ridgway & Godfrey 1997). The Leeuwin Current flows southward
along the Western Australian coast and continues eastward into and across the Great Australian
Bight reaching the west of Tasmania in austral winter (Ridgway & Condie 2004). The influence
of these currents is evident from the occurrence of tropical fauna and flora in southern Australian
waters at normally temperate latitudes (Maxwell & Cresswell 1981, Wells 1985, Dunlop & Wooller
1990, O’Hara & Poore 2000, Griffiths 2003). The importance of these major currents in structuring
marine communities can be seen in the biogeographic distributions of many species, functional
ent
l Curr
Equatoria Coral sea
outh
S South E
quato
Northern r ial Cu
rrent
Territory
Queensland
E as
L e e u w in C
Western
t Australian C
Australia South
Australia
New
ur
ren
South
t
Wales
urre
Great
nt
Victoria
Australian Bight Tasman sea
Tasmania
Figure 2 Major currents and circulation patterns around Australia. The continent is bounded by the Pacific
Ocean to the east, the Indian Ocean to the west and the Southern Ocean to the south. Figure courtesy of
S. Condie/CSIRO.
409
110°00’E 120°00’E 130°00’E 140°00’E 150°00’E

10°00’S N 10°00’S
1b
Darwin
1c?
2a
1a Kimberlays 2c
Cairns
Burketown
20°00’S Port Hedland Northern 20°00’S
Territory Mackey 3
Queensland
Australia
Western Australia
Brisbane
30°00’S South Australia 30°00’S
New South
5a Perth Ceduna Wales
Esperance Sydney
2b Adelaide 2d
Canberra
Victoria ACT
Melbourne 5b
4 2d
40°00’S 40°00’S
Tasmania 2d
6 Hobart
110°00’E 120°00’E 130°00’E 140°00’E 150°00’E
Figure 3 (See also Colour Figure 3 in the insert.) Phytoplankton provinces around Australia. In northern shelf
waters westwards from Torres Strait tropical diatom species dominate, with slight regional differences in
relative abundances and absolute biomass (1a-c). The shallow waters of the Great Barrier Reef region (3) are
dominated by fast-growing nano-sized diatoms. The deeper waters of the Indian Ocean and the Coral Sea are
characterised by a tropical oceanic flora (2a and 2c, respectively) that is dominated by dinoflagellates and
follows the Leeuwin Current (2b) and the East Australia Current and its eddies (2d). South-eastern coastal
waters harbour a temperate phytoplankton flora (4) with seasonal succession of different diatom and dinoflagel-
late communities. Waters south of the tropical and temperate phytoplankton provinces are characterised by
an oceanic transition flora (5a,b) that communicates to the subantarctic phytoplankton province (6) and is
highly variable in extent. The phytoplankton provinces are associated with surface water masses and the
zooplankton fauna likely shows a similar pattern (Figure prepared by G.M. Hallegraeff for CSIRO and National
Oceans Office).
groups and communities. For example, there is broad agreement between phytoplankton community
distributions and water masses (Figure 3).
Australian waters are generally nutrient poor (oligotrophic), particularly with respect to nitrate
and phosphate because the boundary currents are largely of tropical and subtropical origins and
there is little input from terrestrial sources. In general, Australia has a low average annual rainfall
and this rainfall is highly variable. Much of the interior is desert and in the west the aridity extends
to the coast. Monsoonal rains fall in the tropical north during the wet season (December to March)
with cyclones common at this time, but there is little or no rainfall during the rest of the year.
Australian soil is generally low in nutrients and this, together with the high variability in rainfall,
results in little terrestrial nutrient input into the surrounding sea. The generally oligotrophic status
of Australian marine waters contrasts with many mid-latitude productive coastal areas around the
world. This distinction is particularly strong on the western coast of Australia where the Leeuwin
Current replaces the upwelling systems produced by the highly productive eastern boundary currents
characteristic of all other major ocean basins.
The impact of changing productivity on marine oligotrophic systems is largely unknown; they
may not be as resilient to stress and disturbance, including climate change, as more productive
410
systems that commonly experience considerable interannual variability. Changes in the terrestrial
climate also impact Australia’s marine ecosystems to a greater degree than other parts of the world,
so it may not be possible to generalise easily from knowledge elsewhere. Aeolian dust input may
be an important regulator of coastal primary production. In regions south of Tasmania, where
macronutrient concentrations are always high, iron availability influences growth, biomass and
composition of phytoplankton (Sedwick et al. 1999, Boyd et al. 2000). In the macronutrient-limited
regions more typical of the waters around continental Australia, the atmospheric supply of iron
may stimulate nitrogen-fixing phytoplankton, which have a higher iron requirement than other
phytoplankton and therefore influence phytoplankton community composition (Jickells et al. 2005).
Climate-induced changes in wind or rainfall may thus have disproportionately large consequences
for waters around Australia.
Climate change will influence physiology, abundance, distribution and phenology of species
both directly and indirectly, although impacts will usually become most apparent at an ecosystem
level. Given the intrinsic complexity of ecosystems and the uncertainties in future climate projec-
tions, predicting consequences for biodiversity is difficult and highly speculative. Response rates
will depend on the magnitude of changes and on longevity of the species involved in a particular
system. Plankton systems will therefore respond quickly (Hays et al. 2005), whereas a lag might
generally be expected in responses of long-lived species. The ability for adaptation to change will
also vary among species but the rapid rate of present climate change coupled with high exploitation
and destruction or alteration of habitats will compromise the resilience of many populations and
ecosystems (Travis 2002). Strategies for adaptation and mitigation of climate change impacts must
begin with the identification of ecosystems or populations that are most vulnerable to change and
those most vulnerable to other anthropogenic stressors.
In this review, we address the potential impacts of climate variability and climate change on
Australian marine life from the intertidal zone through pelagic waters and into the deep sea. We
provide a synopsis of climate change projections for Australia of key climate variables known to
regulate marine ecosystems from the only IPCC (Intergovernmental Panel of Climate Change)
climate system model constructed in the Southern Hemisphere, the Commonwealth Scientific and
Industrial Research Organisation (CSIRO) Mk3.5 model. Our focus is on the critical variables that
regulate processes in marine ecosystems, namely, temperature, winds, currents, solar radiation,
mixed-layer depth and stratification, pH and calcium carbonate saturation state, storms and precip-
itation, and sea level. We review the expected impacts on species and communities of changes in
each of these variables based on laboratory, modelling and field work and concentrate on biological
groups found in three broad ecosystems: coastal, pelagic and offshore benthic.
Australian marine biodiversity

Australia has highly diverse and unique marine flora and fauna, ranging from spectacular coral
reefs in the tropics to giant kelp forests in Tasmanian waters. The biodiversity of tropical Australia
is high because it is a continuation of the Indo-Pacific biodiversity hot spot, but much of this fauna
is threatened by overharvesting and unregulated development in this region including countries to
the north of Australia. The species diversity of seagrasses and mangroves is among the world’s
highest, particularly in tropical Australia (Walker & Prince 1987, Kirkman 1997, Walker et al.
1999). Temperate Australian waters contain high numbers of endemic organisms due to their long
history of geographic isolation from other temperate regions (Poore 2001). Australian waters also
harbour species and ecosystems that are of international importance. The best-known example is
the Great Barrier Reef, which is the world’s largest World Heritage Area and extends some 2100
km along the coast of northeast Australia.
411
Although Australian temperate waters have lower species diversity than the northern tropical
waters, they harbour much higher numbers of endemic species (Poore 2001). Approximately 85%
of fish species, 90% of echinoderm species and 95% of mollusc species in these southern waters
are endemic (Poore 2001). This high endemism is also documented in Australia’s temperate
macroalgae (Bolton 1996, Phillips 2001). High endemism along the southern coastline is partly
the result of low dispersal abilities of species and the presence of ecological barriers to dispersal
along the southern coastal waters such as a sharp temperature gradient near the cessation of the
Leeuwin Current and the absence of near-shore rocky reefs in the centre of the Great Australian
Bight and at other locations along the southern Australian coastline.
Australia’s fish fauna is extremely diverse and endemic by world standards due to a high
diversity of tropical and temperate habitats and due to the geographic isolation of the temperate
regions. Pelagic fish found around Australia include iconic species such as tuna, billfish (swordfish
and marlin) and sharks. The continental shelf waters off southern Queensland have been identified
as a biodiversity hot-spot for large pelagic fishes (Worm et al. 2003). In contrast to the pattern
elsewhere, this Australian pelagic fish hot spot is located in an area of high catch rates and fishing
effort (Campbell & Hobday 2003). Valuable fisheries exist, despite the generally low productivity
of Australian marine waters; these include the Northern Prawn Fishery, the Southern Bluefin Tuna
Fishery, the Eastern Tuna and Billfish Fishery and the Western Rock Lobster Fishery. Small pelagic
species, such as sardines, jack mackerel, redbait and squid are captured in lower-value but high-
volume coastal fisheries operating from a number of Australian ports. For many of these, there are
well-known correlations between environmental factors and the productivity of the fishery. For
example, the size of the Western Rock Lobster Panulirus cygnus Fishery, which is Australia’s most
important single-species fishery and the world’s largest rock lobster fishery, varies in a predictable
manner with the strength of the Leeuwin Current (Caputi et al. 2001). Similarly, size of banana
prawn Penaeus merguiensis catches in some areas of northern Australia is correlated with wet season
rainfall (Staples et al. 1982, Vance et al. 1985). These variables are likely to change as climate changes.
Further offshore, cold-water corals are found on seamounts and the continental rise, particularly
within the Tasmanian Seamounts Marine Reserve. Cold-water corals are hot spots for biodiversity,
comparable to shallow tropical coral reefs, although little is known of their ecology, population
dynamics or distribution in Australian waters. Over 850 macro- and megafaunal species were recently
found on seamounts in the Tasman and southeast Coral Seas, of which 29–34% were potential
endemics or new to science (Richer de Forges et al. 2000, Williams et al. 2006).
Globally significant populations of many other groups occur in Australia including populations
of marine turtles, marine mammals and seabirds. Six of the seven living species of marine turtle
forage and breed in Australian tropical waters. Marine turtles home to their natal area to breed and
large rookeries used by tens to hundreds of thousands of turtles occur along the northern Australian
coastline and the southern Great Barrier Reef area (Marsh et al. 2001). The flatback turtle Natator
depressus nest only on Australian beaches so can be considered endemic to Australia. The dugong
Dugong dugon forages on seagrasses in tropical Australasian waters. This species is highly threat-
ened in much of its range and a large proportion of global dugong stock is believed to be in Moreton
Bay in eastern Australia and Shark Bay in Western Australia (Marsh et al. 2001). Australian fur
seals Arctocephalus pusillus doriferus, the world’s fourth rarest seal species, and the endemic
Australian sea lion Neophoca cinerea, one of the most endangered pinnipeds in the world, breed
at sites along the southern coast of Australia. These non-migratory pinniped species remain in
southern Australian waters for their entire lives. Around 45 species of whales, dolphins and
porpoises are found in Australian waters including large baleen whales such as the southern right
whale Eubalaena australis and the humpback whale Megaptera novaeangliae, which migrate from
their Southern Ocean feeding grounds to temperate waters around the southern parts of Africa,
South America and Australia and to the tropical waters of the Pacific to breed.
412
A diverse seabird fauna breeds on mainland and island coastlines around Australia; for example
the Houtman Abrolhos Islands on the west coast are an important nesting area for Australian seabirds
in terms of biomass and species diversity (Ross et al. 2001). One of the largest documented colonies
of crested terns Sterna bergii globally (13,000–15,000 nesting pairs) occurs in the Gulf of Carpen-
taria in Australia’s tropical north (Walker 1992). Planktivorous seabirds occur in high numbers in
Australia’s southern temperate waters. For example an estimated 23 million short-tailed shearwaters
Puffinus tenuirostris nest in southeast Australia (Ross et al. 2001).
Climate change projections for Australia

A number of climate models have been used to investigate the response of the ocean-atmosphere
system to increased levels of greenhouse gases and aerosols (Cubasch et al. 2001). This review
examines aspects of climate simulations that are relevant to determining how marine ecosystems
will respond to global climate change. In general, climate model simulations using future greenhouse
gas emission scenarios project oceanic warming, an increase in oceanic stratification and alteration
of mixing depth, changes in circulation, increased pH and rise in sea level, alterations in cloud cover
and ozone levels and thus solar radiation reaching the ocean surface and altered storm and rainfall
regimes (Figure 4). It is very likely that such changes will cause considerable alterations in marine
biological communities (Bopp et al. 2001, Boyd & Doney 2002, Sarmiento et al. 2004).
We use future climate projections over the next century from the CSIRO Mk3.5 climate model
(hereafter called the CSIRO climate model; Appendix 1) using the IS92a future emissions scenario,
often referred to as the ‘business-as-usual’ scenario. Although there are subtle differences between
the CSIRO climate model and other international models, many of the general trends in these fields
are similar and we use the CSIRO climate model to suggest the magnitude of the projected changes
in the set of variables that follow.
HUMAN ACTIVITIES
Increased greenhouse gas Change in UV radiation

concentration levels
Warmer air temperatures
Altered storm Altered atmospheric

regimes/rainfall circulation (winds)
Altered run
o ff
Rise in sea-level
Altered nutrient Altered oceanic
supply and circulation Increased
stratification (currents) dissolved CO2
(mixed layer depth)
Ocean acidification
Warmer sea temperatures
Figure 4 Important physical and chemical changes in the atmosphere and oceans as a result of climate change.
413
Ocean temperature
Waters around Australia are projected to warm by 1–2°C by the 2030s and 2–3°C by the 2070s
(Figure 5). The CSIRO climate model projects the greatest warming off southeast Australia and
this is the area of greatest warming this century in the entire Southern Hemisphere. This Tasman
Sea warming is associated with systematic changes in the surface currents on the east coast of
Australia; including a strengthening of the EAC and increased southward flow as far south as
Tasmania (Figure 5). This feature is present in all IPCC climate model simulations, with only the
magnitude of the change differing among models. Changes in currents leading to the Tasman Sea
warming observed to date is driven by a southward migration of the high-latitude westerly wind
belt south of Australia, and this is expected to continue in the future (Cai et al. 2005, Cai 2006).
10°N 35 10°N 10 10°N 260

240
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0° 0° 0° 220
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25
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180
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20°S 20°S 2 20°S
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0 −6
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60°S −5 60°S −8 60°S 0
60°E 80°E 100°E 120°E 140°E 160°E 180° 60°E 80°E 100°E 120°E 140°E 160°E 180° 60°E 80°E 100°E 120°E 140°E 160°E 180°
20.0 cm/s
10°N 2.6 10°N 6 10°N 10

2.4 5 0
0° 2.2 0° 4 0° −10
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10°S
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10°S
2
10°S −30
−40
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20°S 20°S 20°S −50
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0.8 40°S
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0.6 −4 −100
50°S 0.4 50°S −5 50°S −110
0.2 −6 −120
60°S 0 60°S −7 60°S −130
5.00 cm/s
Figure 5 (See also Colour Figure 5 in the insert.) Simulated annual means of SST (°C) with annual mean
surface currents (cm/s) (left), annual mean zonal winds (m/s) (middle), and mixed layer depth (m) (right). In
the middle panels, westerly wind direction is denoted by positive sign, easterly wind direction by negative
sign. Top row: 1990s, bottom row: difference between 1990s and 2070s.
414
Winds
Under global warming scenarios, the southeasterly trade winds strengthen east of northern Australia,
but weaken to the west of the continent (Figure 5). Westerly winds in southern Australian waters
will weaken. In the Australian coastal region, downwelling will prevail due to the dominating winds
and density structure of the upper ocean. Increasing wind intensity may suppress localised upwelling
in the northeast. However, decreasing wind intensity in southern waters may facilitate localised
upwelling there.
Ocean currents
Surface currents on the east coast will show a systematic change (Figure 5) including EAC
strengthening and increased southward flow as far south as Tasmania. On the west coast there will
be no obvious strengthening of the Leeuwin Current. In the south, the Great Australian Bight region
will experience more westward transport as global temperatures rise. Along the northwest and
northeast coasts there will be an increase in the northward flow.
Mixed-layer depth and stratification

The Australian coastal region is generally a downwelling region due to prevailing winds and density
structure of the ocean. In oligotrophic marine regions of Australia, the dominant mechanism of
nutrient supply to the upper ocean is winter convective mixing due to cooling of surface waters.
Under these conditions the seasonal evolution of the mixed-layer depth and density differences
between this layer and the water below play an important role in the supply of nutrients to the
upper ocean. Surface ocean warming will stabilise the upper ocean and reduce the supply of nutrients
to the surface. The CSIRO climate model simulations project a decline in the annual mean mixed-
layer depth by the 2070s (Figure 5).
CO2, pH and calcium carbonate saturation state

Over the last 200 years, oceans have absorbed 40–50% of the anthropogenic CO2 released into the
atmosphere (Raven et al. 2005). Rising atmospheric CO2 concentrations via fossil fuel emissions
will lead to enhanced oceanic CO2 as the ocean re-equilibrates with the perturbed atmosphere
(McNeil et al. 2003). Elevated CO2 in the upper ocean will alter the chemical speciation of the
oceanic carbon system. As CO2 enters the ocean it undergoes the following equilibrium reactions:
CO 2 + H 2O ⇔ H 2CO3 ⇔ HCO3− + H + ⇔ CO32− + 2H +
Two important parameters of the oceanic carbon system are the pH and the calcium carbonate
(CaCO3) saturation state of sea water (Ω). Ω expresses the stability of the two different forms of
CaCO3 (calcite and aragonite) in sea water.
Increasing CO2 concentration in the surface ocean via uptake of anthropogenic CO2 will have
two effects. First, it decreases the surface ocean carbonate ion concentration (CO32−) and decreases
Ω. Using an ocean-only model forced with atmospheric CO2 projections (IS92a), Kleypas et al.
(1999) predicted a 40% reduction in aragonite saturation (Ωarag) by 2100. Laboratory experiments
415
have shown that some species of corals and calcifying plankton (Gattuso et al. 1998, Langdon et al.
2000, Orr et al. 2005) are highly sensitive to changes in Ω, which has led to the hypothesis of large
decreases in future calcification rates under elevated atmospheric CO2 (Kleypas et al. 1999). Second,
when CO2 dissolves in water it forms a weak acid (H2CO3) that dissociates to bicarbonate, generating
hydrogen ions (H+), which makes the ocean more acidic (pH decreases). Using an ocean-only
model forced with atmospheric CO2 projections (IS92a), Caldeira & Wickett (2003) predicted a
pH drop of 0.4 units by the year 2100 and a further decline of 0.7 by the year 2300. They argued
that the oceanic absorption of anthropogenic CO2 over the next several centuries may result in a
pH decrease greater than inferred from the geological record over the past 300 million years, with
the possible exception of those resulting from rare, extreme events such as meteor impacts.
Changes in surface pH and in Ωarag reflect changes in the speciation of carbon within the ocean
and are a function of temperature, salinity, alkalinity and dissolved inorganic carbon concentrations.
McNeil & Matear (2006) showed that climate change does not alter the projected change in surface
pH. The projected pH decrease is controlled by the future levels of atmospheric CO2. However,
the decline in Ωarag due to rising CO2 levels in the ocean is slightly reduced (~15%) because of the
increase in Ωarag due to the increase in surface temperature. For the Australian region, the pH and
Ωarag for the 1990s are shown along with the corresponding change in these values relative to 1990s
(Figure 6). We see significant declines in these parameters but with the greatest declines occurring
off northeast Australia. A major unknown in this region is whether any dissolution of the tropical
coral reefs would buffer the pH decreases. Because of the enhanced levels of CO2 in the atmosphere
and rates of fossil fuel burning, the process of ocean acidification is essentially irreversible over
the next century. It will take thousands of years for ocean chemistry to return to a condition similar
to that of preindustrial times.
Solar radiation
Highly energetic ultraviolet radiation (UVR) penetrates the ocean surface and is known to have
detrimental effects on marine organisms. UVR penetration to the earth’s surface increased during
the last quarter of the twentieth century as stratospheric ozone was depleted by chlorofluorocarbons
(CFCs), halons, hydrochlorofluorocarbons and other compounds. Stratospheric ozone levels appear
to have stabilised, however, due to the 1989 implementation of the Montreal Protocol designed to
phase out the production of CFCs and other compounds that deplete the ozone layer (de Jager et al.
2005).
Most climate models predict that the ozone layer will recover and thicken throughout the
twenty-first century (de Jager et al. 2005), so UVR penetration should decline (McKenzie et al.
2003). However, these predictions are somewhat uncertain, especially in the timing of the rethick-
ening, due to uncertainties in projections of greenhouse gas emissions and degradation and due to
the complex ways that chemical, radiative and dynamic processes will affect stratospheric ozone.
For example, chemical reactions of some greenhouse gases (such as methane) can reduce total
ozone in the stratosphere but the level of methane emissions is difficult to predict. Climate change
will also affect UVR penetration indirectly by influencing other factors such as aerosols, clouds
and snow cover. Aerosols can scatter more than 50% of the UV-B — the biologically important
component of UVR — and aerosols increased in the atmosphere during most of the twentieth
century, although they have shown declines since 1990 (Schiermeier 2005). Clouds can attenuate
15–30% of the UV-B, and cloud reflectance measured by satellite has shown a long-term increase in
some regions of the world (McKenzie et al. 2003). All these factors introduce considerable uncer-
tainty in future levels of UVR at the ocean surface, and it has been suggested that climate warming
will slow the recovery of the ozone layer by up to 20 yr (Kelfkens et al. 2002).
416
10ºN 8.16 10ºN 7

8.14 6.5
0º 0º
8.12
6
10ºS 8.1 10ºS
5.5
8.08
20ºS 20ºS 5
8.06
30ºS 8.04 30ºS 4.5

8.02
4
40ºS 8 40ºS
3.5
7.98
50ºS 50ºS
7.96 3
60ºS 7.94 60ºS 2.5

60
80
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12
14
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18
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80
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18
ºE
ºE
0º
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ºE
ºE
0º
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0º
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0º
E
E
10ºN −0.09 10ºN −0.5
−0.1 −0.6
0º 0º
−0.11 −0.7
10ºS −0.12 10ºS −0.8
−0.13 −0.9
20ºS 20ºS
−0.14 −1
30ºS −0.15 30ºS −1.1
40ºS −0.16 40ºS −1.2

−0.17 −1.3
50ºS 50ºS
−0.18 −1.4
60ºS −0.19 60ºS −1.5

60
80
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80
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ºE
ºE
0º
0º
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0º
0º
ºE
ºE
0º
0º
0º
0º
0º
E
Figure 6 (See also Colour Figure 6 in the insert.) Simulated annual means of pH (left) and aragonite saturation
state (right). Top row: 1990s, bottom row: difference between 1990s and 2070s.
Precipitation and storms

Changes in the amount or timing of rainfall and the associated river runoff affect the salinity regimes
of estuaries and adjacent coastal waters, while in comparison salinity is relatively constant through-
out the year in most oceanic waters. Despite the high uncertainty of rainfall projections in Australia,
there is a tendency for decreased rainfall over most of Australia and over the oceans in climate
model simulations (Figure 7). This general reduction in rainfall may be offset by an increase in
the frequency of intense storms (Emanuel 2005, Webster et al. 2005), which will increase rainfall
intensity and the associated runoff of freshwater and suspended sediments. In northern Australia,
tropical cyclones are important extreme rainfall events. A recent study under 3 times the baseline
levels of CO2 conditions based on levels prior to the industrial revolution in the mid-1800s, projected
a 56% increase in the number of simulated tropical cyclones over northeastern Australia with peak
winds greater than 30 ms−1 (Walsh et al. 2004). However, the behaviour of tropical cyclones under
417
10ºN 280 10ºN 15 10ºN 260

260 14 240
0º 0º 0º 220
240 12
10 200
10ºS 220 10ºS 10ºS
180
8
200 20ºS 160
20ºS 6 20ºS
180 140
30ºS 4
30ºS 30ºS 120
160
2
100
40ºS 140 40ºS 0 40ºS
80
120 −2
50ºS 60
50ºS 50ºS
100 −4 40
60ºS 80 60ºS −6 60ºS 20
60
80
10
12
14
16
18
60
80
10
12
14
16
18
60
80
10
12
14
16
18
ºE
ºE
0º
0º
0º
0º
0º
ºE
ºE
0º
0º
0º
0º
0º
ºE
ºE
0º
0º
0º
0º
0º
E
E
E
E
E
E
E
E
E
E
E
E
10ºN 100 10ºN 3 10ºN 28
2.5 26
80
0º 0º 0º 24
2
60 22
10ºS 10ºS 1.5 10ºS 20
40
1 18
20ºS 20 20ºS 20ºS
0.5 16
0 0 14
30ºS 30ºS 30ºS
−0.5 12
−20 10
40ºS 40ºS −1 40ºS
−40 8
−1.5
50ºS 50ºS 50ºS 6
−60 −2 4
60ºS −80 60ºS −2.5 60ºS 2
60
80
10
12
14
16
18
60
80
10
12
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18
60
80
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º
ºE
0º
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ºE
ºE
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ºE
ºE
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0º
0º
E
E
E
E
E
E
E
E
E
E
E
E
E
Figure 7 (See also Colour Figure 7 in the insert.) Simulated annual means of downward solar radiation at
the ocean surface (W/m2) (left), precipitation minus evaporation (mm/d) (middle), and sea-level height anomaly
due to upper ocean stratification relative to 2000 m (cm) (right). Top row: 1990s, bottom row: difference
between 1990s and 2070s.
global warming is uncertain because they are not currently well resolved by global or regional
climate models (Pittock et al. 1996, Walsh & Pittock 1998).
Sea level
Rising sea level around Australia will flood existing coastal environments and alter their marine
habitats. With global warming, the CSIRO climate model projects a doubling in the rate of sea-
level rise from the observed 1.44 mm yr−1 for the twentieth century (Church et al. 2001). By the
2080s, sea level is projected to rise by 0.06–0.74 m above the 1990 value (Gregory et al. 2001).
These projections take into account both the mean global projections from the IPCC scenarios and
the non-uniform spatial distributions of sea-level change related to thermal expansion produced by
the climate simulations. However, they do not include vertical land movement, which can be locally
important. Sea-level rise projected by the CSIRO model for just the thermal expansion shows an
increase in the entire Australian region but with large spatial variability (Figure 7). The variability
in sea-level rise reflects how the excess heating of the planet due to global warming is stored in
418
the oceans, and this large variability is supported by reconstructed sea-level estimates from the past
decade (Willis et al. 2003). Therefore, over this century the local impact of sea-level rise may
substantially deviate from the global averaged value. For the Australian region, much greater sea-
level rise is projected on the east coast than the west coast due to the increased southward penetration
of the warm EAC, which causes water here to expand more than in other regions.
Climate impacts on Australian marine life

In this section we describe the impacts of climate variables on marine life in coastal, pelagic and
offshore benthic systems. We consider the climate variables that have greatest impact on structuring
marine communities within these systems and for which projections over the next 100 yr are
available from global climate models. Where applicable, we review impacts on physiology, distri-
butions and abundance, and phenology of marine organisms. Studies of climate impacts from both
field and experimental research from Australia are discussed and supplemented with studies and
observations from international research. Results of this section are summarised in Table 1.
Ocean temperature
Elevated water temperatures stress plants and animals already near the upper limits of their optimal
temperature range, slowing growth and impairing reproductive capacity (Philippart et al. 2003,
Roessig et al. 2004, Helmuth et al. 2005, Keser et al. 2005). This is because most biological
processes have an optimal temperature range and outside this range physiological efficiency
declines.
Coastal systems
Physiology Extreme temperatures, both warm and cool, if severe or prolonged can lead to irrep-
arable damage and death of coastal organisms as well as photosynthetic inhibition in marine plants
(Bruhn & Gerard 1996, Ralph 1998, Davenport & Davenport 2005, Campbell et al. 2006). Large
diebacks of marine fauna and flora in the intertidal and shallow subtidal occur on very hot days
particularly when these coincide with low tides during the middle of the day (Tsuchiya 1983, Perez
et al. 2000). Such a situation may have been responsible for the major dieback of seagrass beds in
southern Australia during early 1993 when over 12,000 hectares were lost (Seddon et al. 2000).
Probably the most widely publicised mass mortalities induced by warmer-than-average tem-
peratures are those resulting from tropical coral reef bleaching events (Hoegh-Guldberg 1999).
During bleaching events, the symbiosis between the coral and the unicellular algae (dineflagellates
from the genus Symbiodium) that live within the coral tissues disintegrates. Bleached corals may
recover their symbiotic populations of Symbiodium in the weeks and months after a bleaching event
if the conditions triggering the event are mild and short-lived, but mortality has reached 100% in
bleached corals when stressful conditions have persisted for days to weeks. Recent warming
throughout tropical oceans has led to repeated coral bleaching events, not seen anywhere in the
world before 1979, affecting hundreds to thousands of square kilometres of coral reefs in almost
every region of the world where coral reefs occur. In the most severe global episode of mass coral
bleaching (1998), 16% of corals that were surveyed before that event had died by the end of the
year (Hoegh-Guldberg 1999, Knowlton 2001).
Mass bleaching events over large sections of the Great Barrier Reef have occurred six times
during the past 30 years: in 1983, 1987, 1991, 1998, 2002 and 2006. Mortality rates in this region
were relatively low however, primarily because warming on the Great Barrier Reef was less severe
than in other parts of Australia and the world. For example, in 1998 a very warm pool of water sat
419
Table 1 Expected and observed impacts of climate change on Australian marine life and field
or experimental evidence from outside Australia
Observations
elsewhere or
Expected change Species group/ Expected climate impact Observations experimental
in climate natural system in Australia in Australia evidence
Increasing Seagrasses Poleward shift in species ranges Seagrass
temperature and and a shift in abundance toward distributional limits
mangroves species tolerant of warmer linked to
waters temperature1
Earlier flowering and fruiting Flowering of
seagrasses in
temperate Australia
linked to water
temperature2
Seagrass Increased frequency and Southern Australia
intensity of large-scale early 1993
diebacks with increase in (>12,000 hectares)3
frequency and intensity of
extreme temperatures
Rocky shore, Poleward shift in species ranges Rocky shores in
fauna and and a shift in abundance toward Europe, United
macroalgae species tolerant of warmer States and South
waters America over past
50 yr4
Increased frequency and Diebacks in Tasmania European and
intensity of large-scale and South Japanese coasts6
diebacks with increase in Australian hot days5
frequency and intensity of
extreme temperatures
Kelp Contraction of kelp ranges, Decline of kelp in Loss of kelp in east
communities declines in abundance, local Tasmanian waters Pacific following
extinctions, particularly in over past 50 yr7 El Niño8
Tasmania
Phytoplankton Poleward shift in species ranges Southward extension Poleward shift in
and a shift in abundance toward of a coccolithophore North Atlantic10
warm-water species and a dinoflagellate
in southeast
Australia9
A decline where warming North Atlantic11
enhances stratification
Earlier appearance of plankton North Sea12
in summer in temperate waters
Increase in frequency and Norwegian coast13
intensity of harmful and
nuisance blooms
Zooplankton Poleward shift in species ranges Large poleward
and a shift in abundance toward range shifts
warm-water species (>1000 km) in
North Atlantic14
A decline where warming North Atlantic15
enhances stratification
420
Table 1 (continued) Expected and observed impacts of climate change on Australian marine
life and field or experimental evidence from outside Australia
Observations
elsewhere or
Earlier appearance of North Sea16
zooplankton in summer in
temperate waters
Coral reefs Increase in frequency and Six severe bleaching Coral reefs
severity of coral bleaching and events in past 30 yr globally18
mortality (Great Barrier Reef,
Ningaloo Reef)17
Increase in local extinctions of Coral reefs
coral-associated fauna with globally19
bleaching events
Demersal and Poleward shift in species ranges Tasmanian fish North Atlantic fish
pelagic fish and a shift in abundance toward distributions shifting
species tolerant of warmer shifting south with northward21
waters increase in fish that
prefer warmer
waters20
Earlier dates of mean migration Earlier migrations
and spawning in temperate and in northeast
subtropical species Atlantic fish22
Seabirds and Poleward shifts in species Southward shift of
wetland birds ranges and a shift in abundance seabird distributions
toward species tolerant of in Western Australia
warmer waters and increase in
abundance23
Earlier arrival in migratory Southern Australian Terrestrial, wetland
species in temperate and wetland birds24 and seabirds
subtropical regions globally25
Earlier nesting and laying and Western and southern
protracted breeding seasons in Australian
temperate and subtropical seabirds26
species
Marine turtles Poleward shift in species Northward shift of
and foraging ranges cetaceans and
mammals turtles in northeast
Atlantic27
Earlier breeding Earlier nesting in
marine turtles in
United States28
Skewing of turtle sex ratios Experimental and
toward females modelling
evidence that
warmer
temperatures
produce more
females29
421
Observations
elsewhere or
Alteration of Phyto- and Increased productivity where Production pulses Decreased
winds zooplankton wind mixing is enhanced and a correlated with production in
reduction where wind strength peaks in wind central North
declines oscillation in Pacific during
Tasmanian shelf low-wind
waters30 regimes31
Coastal fish Recruitment strength linked to Rocky reef fish32
wind strength
Seabirds Reduction of breeding success Breeding colonies on
with prolonged periods of Great Barrier Reef33
strong winds
Alteration of Seagrasses & Local extinctions of cold-water Seagrass
currents mangroves species in southeastern distributional limits
including Australia with increased flow further south on
strengthening of EAC, appearance of tropical west coast than east
of EAC species further south on east coast due to
coast influence of warm-
water Leeuwin
Current34
Rocky shore, Local extinctions of cold-water Tropical species
fauna and species in southeastern already found at
macroalgae Australia with increased flow temperate latitudes
of EAC, appearance of tropical on east coast35
species further south on east
coast
Kelp Local extinctions of cold-water Expansion of long-
communities species in southeastern spined urchin to
Australia with increased flow Tasmania facilitated
of EAC, appearance of tropical by larval transport
species further south on east by EAC36
coast
Phyto- and Poleward extension of warm High abundance of a
zooplankton currents will transport tropical tropical
plankton more southward coccolithophore off
southeast Australia37
Decline in Phyto- and Decrease in abundance Phytoplankton
mixed-layer zooplankton productivity in
depth/increasing central North
stratification Pacific declines as
mixed-layer depth
decreases38
Increased CO2 and Mangroves Increase in productivity with Experimental
decrease in pH rising atmospheric CO2 evidence39
and aragonite
saturation state
422
Observations
elsewhere or
Seagrasses Increase in productivity with Experimental
increase dissolved CO2 and evidence40
deepening of depth limits
Rocky shore, Impaired growth in calcifying Experimental
fauna and fauna and macroalgae and evidence41
macroalgae increase in mortality of early
life stages
Phytoplankton Changes in growth and Experimental
community composition; long- evidence42
term decline in abundance and
distribution of calcifying
species
Zooplankton Impaired growth in calcifying Experimental
species, particularly pteropods; evidence43
midterm decline in abundance
and distribution
Coral reefs Impaired growth rates and Experimental and
possible dissolution modelling
evidence44
Cold-water High threat of impaired growth Evidence from
corals rates and possible dissolution modelling work45
Possible increase Seagrasses Reduction of growth rates and Experimental
in UV biomass in UV-sensitive evidence46
species
Mangroves Reduction of growth rates and Experimental
biomass in UV-sensitive evidence47
species
Rocky shore Increase mortality of early life Experimental
fauna and stages and reduction of growth evidence48
macroalgae rates in UV-sensitive species
Kelp and Increase mortality of early life Experimental
subtidal stages evidence49
macroalgae
Phytoplankton Reduction of growth rates and Evidence from field
biomass in UV-sensitive and laboratory
species and of nutritional value experiments50
to zooplankton
Changes in community
composition
Zooplankton Increased mortality of early life Evidence from
stages and reduction of growth laboratory
rates in UV-sensitive species experiments51
Coral reefs Increase in mortality during Evidence from
bleaching events through syn- laboratory
ergistic effects with temperature experiments52
423
Observations
elsewhere or
Increase mortality of early life Evidence from
stages and reduction of growth laboratory
rates experiments53
Demersal and Damage to epidermis and ocular Evidence from
pelagic fish components in pelagic species laboratory
and increased mortality in egg experiments54
and larval stages in shallow
water and upper ocean
Increase in Mangroves Shifts in community abundance Increase in mangrove
frequency or as coastal salinity regimes are area in southeast
intensity of altered and nutrient and Australia may be
severe storms sediment loading changes indirectly linked to
and extreme changes in rainfall
rainfall events although changes in
and a decrease in land use likely to be
average rainfall overriding factor55
Seagrasses Destruction of seagrass beds Loss of >1000 km2 in Large-scale
Harvey Bay after destruction in
severe storms and United States after
flooding56 cyclones57
Kelp Shifts in community abundance Switch from canopy- Range shifts of
communities and increased local mass forming macroalgae macroalgae in
and subtidal mortality events associated to turf-forming algae New Zealand and
macroalgae with storms and flood events in South Australia California
linked to enhanced associated with
nutrient supply from storms and wave
coastal runoff58 exposure59
Benthic Shifts in community abundance Mass mortality of Field experiments
macrofauna and increased local mass grazing urchins after revealed shift in
mortality events associated freshwater pulse60 community
with storms and flood events composition with
increased
sedimentation61
Alteration of peak timing of life High rainfall may High rainfall may
cycle events decrease salinity in decrease salinity
estuaries so in estuaries so
triggering prawn triggering prawn
emigration in emigration in the
northern Australia62 United States63
Coral reefs Mass mortality events Mass mortality of Mass mortality of
associated with storms and corals on Great corals in
flood events Barrier Reef after Caribbean after
cyclones and flood cyclones65
events64
424
Observations
elsewhere or
Community structure influenced Lower coral diversity
by rainfall regime and runoff on Great Barrier
Reef in wet tropics66
Phytoplankton Diatoms may decline with Evidence from field
decreasing average runoff and experiment and
nutrient input while dino- time series67
flagellates (including harmful
algae) may profit from storm-
associated runoff and humic
substances in coastal waters
Marine turtles Increased mortality events High mortalities of
and turtles and seal pups
mammals associated with
cyclones and
storms68
Rise in sea level Mangroves Alteration of hydrological or Mangroves in
tidal regimes leads to mortality Africa and Asia69
of mangroves
Mangrove retreat with rising sea Caribbean70
level
Seagrass Reduction in growth of seagrass 50 cm rise in sea
and distributional shifts level expected to
result in 30–40%
reduction of
seagrass growth71
Seabirds Loss of breeding sites for Evidence from
species that nest on low-lying modelling work72
coastal areas through increased
flooding and erosion
Marine turtles Loss of breeding and haul-out 50 cm rise in sea
and sites for species through level expected to
mammals increased flooding and erosion lead to a 32% loss
of turtle nesting
beaches in the
Caribbean73
Notes: 1Walker & Prince 1987; 2West & Larkum 1979, Cambridge & Hocking 1997, Inglis & Smith 1998; 3Seddon et al.
2000; 4Barry et al. 1995, Southward et al. 1995, Sagarin et al. 1999, Zacherl et al. 2003, Mieszkowska et al. 2005, Rivadeneira
& Fernandez 2005, Simkanin et al. 2005, Smith et al. 2006; 5Valentine & Johnson 2004, Womersley & Edwards 1958;
6Tsuchiya 1983, Perez et al. 2000; 7Edyvane 2003, Edgar et al. 2005; 8Dayton & Tegner 1984, Zimmerman & Robertson 1985,
Dayton et al. 1998, 1999, Adey & Steneck 2001; 9Blackburn & Creswell 1993, Blackburn 2005, G. Hallegraef pers. com.;
10M. Edwards 2005; 11Richardson & Schoeman 2004; 12Edwards & Richardson 2004; 13Edwards et al. 2006; 14Beaugrand
et al. 2002, Bonnet et al. 2005; 15Richardson & Schoeman 2004; 16Greve et al. 2004, Edwards & Richardson 2004, Kirby
et al. 2007; 17Hoegh-Guldberg 1999, Wilkinson 2004; 18Hoegh-Guldberg 1999, Knowlton 2001; 19Dulvy et al. 2003;
20Welsford & Lyle 2003, P. Last pers. com.; 21Beare et al. 2004, Byrkjedal et al. 2004, Perry et al. 2005, (continued on next page)
425
Notes (continued): Rose 2005a, 2005b; 22Sims et al. 2001; 23Dunlop & Wooller 1986, Dunlop et al. 2001, Bancroft et al.
2004; 24Beaumont et al. 2006; 25Mason 1995, Crick et al. 1997, Archaux 2003, Both et al. 2004, Lehikoinen et al. 2004,
Both et al. 2005, Marra et al. 2005, Jonzén et al. 2006, Moller et al. 2006; 26Dunlop & Wooller 1986, Chambers 2004;
27Robinson et al. 2005, MacLeod et al. 2005, McMahon & Hays 2006; 28Weishampel et al. 2004; 29Yntema & Mrosovsky
1982, Godfrey et al. 1999, Booth & Astill 2001, Glen & Mrosovsky 2004; 30Harris et al. 1991; 31Polovina et al. 1994;
32Thresher et al. 1989; 33King et al. 1992; 34Walker & Prince 1987; 35Griffiths 2003; 36Johnson et al. 2005; 37Blackburn &
Cresswell 1993, Blackburn 2005; 38Venrick et al. 1987, Polovina et al. 1994, 1995; 39Polovina et al. 1995, Roemmich &
McGowan 1995, Farnsworth et al. 1996, Ainsworth & Long 2005; 40Invers et al. 1997, 2002, Zimmerman et al. 1997;
41Gao et al. 1993, Kurihara et al. 2004, Michaelidis et al. 2005, Berge et al. 2006; 42Riebesell et al. 2000, Antia et al. 2001,
Tortell et al. 2002, Engel et al. 2005; 43Orr et al. 2005; 44See Hoegh-Guldberg 2004; 45Guinotte et al. 2006, Raven et al.
2005; 46Dawson & Dennison 1996; 47Moorthy & Kathiresan 1997, 1998; 48Graham 1996, Rijstenbil et al. 2000, Cordi
et al. 2001, Lesser et al. 2003, Przeslawski et al. 2004, 2005, Bonaventura et al. 2006; 49Graham 1996, Bischof et al. 1998,
Swanson & Druehl 2000, Wiencke et al. 2006; 50Behrenfeld et al. 1993, Keller et al. 1997, Wilhelm et al. 1997, Wängberg
et al. 1999, Garde & Cailliau 2000, Barbieri et al. 2002, Litchman & Neale 2005; 51Karanas et al. 1979, Damkaer & Dey
1983; 52Lesser 1996, 1997, Baruch et al. 2005, Drohan et al. 2005; 53Shick et al. 1996, Wellington & Fitt 2003; 54Hunter
et al. 1982, Keller et al. 1997, Zagarese & Williamson 2001, Markkula et al. 2005; 55Saintilan & Williams 1999, Harty
2004, Rogers et al. 2006; 56Preen et al. 1995; 57Thomas et al. 1961; 58Gorgula & Connell 2004; 59Graham 1997, Cole et al.
2001; 60Andrew 1991; 61Norkko et al. 2002, Thrush et al. 2003a, 2003b, Lohrer et al. 2004; 62Staples 1980, Vance et al.
1985, Staples & Vance 1986, Vance et al. 1998; 63Zein-Eldin & Renaud 1986; 64Alongi & Robertson 1995, Alongi &
MacKinnon 2005; 65Porter & Meier 1992, Gardner et al. 2005; 66De Vantier et al. 2006; 67Carlsson et al. 1995, Goffart
et al. 2002; 68Limpus & Reed 1985, Pemberton & Gale 2004; 69Blasco et al. 1996; 70Ellison 1993, Parkinson et al. 1994;
71Short & Neckles 1999; 72Galbraith et al. 2002, Smart & Gill 2003; 73Fish et al. 2005.
above Scott Reef off northwest Australia for several months, resulting in an almost total bleaching
of these offshore reefs and mortality of corals down to 30 m depth. The recovery of Scott Reef
has been very slow (Wilkinson 2004).
By the middle of this century, temperature thresholds for coral bleaching will be exceeded
every year in Australia if sea temperatures increase as projected by global climate models (Hoegh-
Guldberg 1999). Based on the current responses of corals, it is estimated that an increase of 2°C
in tropical and subtropical Australia would result in annual bleaching and quite possibly regular,
large-scale mortalities (Hoegh-Guldberg 1999, 2004, Lough 2000). A geographic analysis of risk
to the Great Barrier Reef associated with these changes in sea temperature indicated that the
projected succession of devastating mass coral bleaching events will severely compromise the
ability of reefs to recover, no matter where they are found along the Queensland coastline (Done
et al. 2003). This analysis indicated that deterioration of coral populations is likely in most of the
scenarios examined and this is reinforced by findings from other studies (Hoegh-Guldberg 1999,
Donner et al. 2005).
For large, mobile animals that may be transient visitors to coastal waters, oceanic warming
may impact particular life stages such as juveniles or embryos. For example, gender in all turtles
is determined by ambient nest temperatures during embryonic development (Mrosovsky et al. 1992,
Godfrey et al. 1999, Hewavisenthi & Parmenter 2002a). Small changes in temperature close to the
pivotal temperature at which a 50:50 sex ratio is produced (~29°C for marine turtles) skew the sex
ratio of hatchlings, with warmer temperatures producing more females (Yntema & Mrosovsky
1982, Godfrey et al. 1999, Booth & Astill 2001, Glen & Mrosovsky 2004). Many nesting beaches
around the world, including most Australian beaches, already have a strong female bias (Limpus
426
1992, Loop et al. 1995, Godfrey et al. 1996, Binckley et al. 1998, Hewavisenthi & Parmenter 2002b,
Hays et al. 2003, Glen & Mrosovsky 2004) so if temperatures rise, the proportion of eggs developing
as males may be further reduced. However, light-coloured (thus cooler) beaches within nesting
regions produce more males (Hays et al. 2003). In Queensland beaches on offshore coral cays and
islands have lighter-coloured sand than mainland beaches, thus maintaining sex ratios (Environment
Australia 1998). Therefore if temperatures warm on these beaches, the gross skewing in sex bias
may have serious implications for local breeding population persistence.
On a global scale outbreaks of disease have increased over the last three decades in many
marine groups including corals, echinoderms, mammals, molluscs and turtles (Ward & Lafferty
2004). Causes for increases in diseases of many groups remain uncertain, although temperature is
one factor that has been implicated in corals, molluscs and turtles (Harvell et al. 2002). Previously
unseen diseases have also emerged in new areas through shifts in distribution of hosts or pathogens,
many of these shifts are in response to climate change (Harvell et al. 1999). A consequence of
climate-mediated physiological stress is that host resistance to pathogens or parasites can be
compromised (Scheibling & Hennigar 1997, Garrabou et al. 2001, Lee et al. 2001, Harvell et al.
2002, Mouritsen et al. 2005). Temperature-induced disease outbreaks in corals on the Great Barrier
Reef have occurred at the same time as bleaching events, resulting in increased coral mortality
rates (Jones et al. 2004). A large-scale mortality of greenlip abalone, Haliotis laevigata, along the
south Australian coast in 1985 and 1986 due to infection by Perkinsus parasites may have been
aggravated by warmer water temperatures predisposing the abalone to this disease (Goggin & Lester
1995). Population declines due to temperature-related disease susceptibility have also been reported
in several Californian abalone species through both observational and experimental studies (Davis
et al. 1996, Vilchis et al. 2005).
Fibropapillomatosis, a disease that causes tumours, is now common in green turtles Chelonia
mydas and olive ridley turtles Lepidochelys olivacea (Adnyana et al. 1997, Jones 2004). This disease
was first documented in the 1930s and was rare until the early 1980s but has since reached epidemic
proportions in many turtle populations worldwide (Jones 2004). The prevalence of the tumours in
young turtles suggests prolonged exposure to anthropogenic pollutants may be responsible
(Adnyana et al. 1997, Herbst et al. 2004, Jones 2004, Ene et al. 2005, Foley et al. 2005). However,
the increase of this disease in recent decades coincides with rapidly rising temperatures so it may
also be indirectly related to climate change (Robinson et al. 2005).
Distribution and abundance Temperature influences the abundance and distribution of coastal
marine life such as macroalgae, seagrasses and molluscs (McMillan 1984, Walker & Prince 1987,
Jernakoff et al. 1996, Steneck et al. 2002, Hiscock et al. 2004). Fluctuations in species abundances
and community composition have been linked to variations in temperature (Southward et al. 1995,
Tegner et al. 1996, Dayton et al. 1999, Grove et al. 2002, M.S. Edwards 2004, Schiel et al. 2004,
Smith et al. 2006). Shifts in species distributions associated with ocean warming are documented
from rocky shores in Europe, the United States and South America (Barry et al. 1995, Sagarin et al.
1999, Zacherl et al. 2003, Mieszkowska et al. 2005, Rivadeneira & Fernandez 2005, Simkanin et al.
2005). For example, a recent comprehensive resurvey of rocky intertidal shores around the United
Kingdom found range extensions in the northern (high-latitude) limits of some warm-water species
over the past 50 yr and a retraction in the southern limits of fewer cold-water species although
rates of recession were not as fast as rates of advancement in warm-water species (Mieszkowska
et al. 2005). The high levels of endemism along Australia’s southern coastline could increase
vulnerability to temperature increases compared to temperate rocky shores elsewhere; many
endemic species may have more stringent temperature limits and so may be particularly susceptible
to warming (Beardall et al. 1998).
427
There are interactive effects between the impacts of warming and availability of nutrients on
distribution and abundance of macroalgae. Declines of giant kelp forest communities in Tasmanian
coastal waters have been associated with thermal and nutrient stress (Edyvane 2003, Edgar et al.
2005). Macrocystis kelp forests in Australia are found predominantly in the southeast where water
conditions are cool and relatively nutrient rich. There has been a considerable decline in Tasmanian
kelp forests over the past 50 yr associated with rising temperatures (Edyvane 2003). Further, an
unusual dieback of the shallow sublittoral brown macroalga Phyllospora comosa along the east
coast of Tasmania in 2001 has also been attributed to above-average seawater temperatures coupled
with nutrient stress (Valentine & Johnson 2004). If the EAC strengthens as projected by climate
models, warm, nutrient-poor water will impinge more frequently on Tasmanian giant kelp commu-
nities, potentially leading to local extinction and a shift of macroalgal communities to understorey-
dominated forms (Kennelly 1987a,b, Dayton et al. 1999).
Globally, mangrove distribution is generally constrained by the 20°C winter sea isotherm; there
are a few exceptions, such as the more southerly distribution of mangroves in eastern Australia
(Duke 1992). It has been suggested that this distribution is the result of small-scale extensions of
warmer currents, such as the EAC, or that the southern populations are a relict representing refuges
of more poleward distributions in the past (Duke 1992). As mangrove species show considerable
variation in their sensitivity to temperature, species composition of mangrove forests will alter as
temperatures rise and species distributions are expected to shift poleward (Field 1995).
Evidence suggests that some benthic and demersal fish species may be able to move as oceans
warm, regardless of whether there is a shift in associated habitats such as coral reefs, kelp forests
or rocky reef communities. Certain fishes associated with coral reefs appear to be able to populate
reefs that do not have corals, as shown by the appearance of coral reef fishes in southern New
South Wales and Victoria during the summer (Hoegh-Guldberg 2004). These fishes recruit into
coastal areas and grow for several months, disappearing when cold conditions return. Many coral
reef fish may be able to move southward as oceans warm, although obligate corallivorous species
would presumably be missing (Hoegh-Guldberg 2004). This has already been observed in other
parts of the world such as California, where the composition of near-shore rocky reef fish commu-
nities shifted in dominance from cold-water northern species to warm-water southern species as
temperatures warmed (Holbrook et al. 1997). However, coral bleaching has already led to local
extinctions of a few coral-associated fish (Dulvy et al. 2003) and doubtless many more could
disappear as coral bleaching episodes increase.
Other mobile groups such as seabirds and marine mammals may be able to rapidly shift their
distributions with climate change, although many are restricted to coastal habitats during breeding
seasons. Warmer waters may allow marine turtles and dugongs to extend their foraging distributions
in Australian inshore waters further south. However, green turtles Chelonia mydas and dugongs
Dugong dugon selectively feed on seagrasses while hawksbill turtles Eretmochelys imbricata forage
on coral reefs, so their ability to shift distributions are likely to be limited by changes in the
distribution of their food sources.
Range expansions have already been observed in seabird species along the west coast of
Australia, with tropical species extending their breeding and foraging ranges southward (Dunlop &
Wooller 1986, Dunlop et al. 2001). The recent growth of nesting colonies of wedge-tailed shear-
waters Puffinus pacificus in southwestern Australia may be due to a southerly movement from more
northerly colonies as temperatures rise (Bancroft et al. 2004). Wedge-tailed shearwaters are found
only over waters with surface temperatures exceeding 20°C (Surman & Wooller 2000). The pop-
ulation of Australasian gannets Morus serrator that breed in southeast Australia has increased by
approximately 6% per year since 1980, with new breeding sites being established as nesting space
becomes limited (Bunce et al. 2002). This increase appears to be associated with a long-term
428
warming trend and a concurrent increase in the abundance of small pelagic prey fish, principally
pilchards Sardinops sagax.
Phenology Water temperature and day length are the principal triggers or correlates for the timing
of biological events such as breeding or migration in marine animals and flowering and seed
germination in marine plants (Parmesan & Yohe 2003). Synchrony in reproduction of widely
distributed seagrass beds and mangroves (Clarke & Myerscough 1991, Inglis & Smith 1998, Diaz-
Almela et al. 2006) suggests control by these environmental variables. Such synchronies of bio-
logical events in distant populations may be regulated by a large-scale independent factor such as
temperature or day length. Regular flowering of the seagrass Posidonia australis occurs between
April and June in southwestern Australia, probably induced by a seasonal decline in water temper-
atures (West & Larkum 1979, Cambridge & Hocking 1997). However, further north in Shark Bay
P. australis meadows do not flower every year (Larkum 1976). Widespread flowering P. australis
is also rare off central New South Wales on the east coast (Walker et al. 1988). Shark Bay and
central New South Wales are near the northern limits for this temperate seagrass species so the
threshold decline in water temperature required to trigger flowering may begin to occur less
frequently. As a warming of coastal waters is projected, particularly off southeast Australia, episodes
of flowering of P. australis may become even rarer in northern meadows. The deposition of seed
banks after flowering is an important process that allows seagrass beds to recover rapidly from
catastrophic disturbances such as storms or floods (Preen et al. 1995).
Temperature has also been correlated with the timing of mass spawning in tropical reef corals
on the Great Barrier Reef (Babcock et al. 1986) and on the tropical west coast (Simpson 1991).
However, the physiological and evolutionary mechanisms that underlie the timing of reproduction
in corals and in most marine invertebrates are far from clear; thus it is difficult to speculate on the
consequences of any change in the timing of spawning.
There is global evidence that climate change is influencing the phenology of larger marine
fauna. Marine turtles in Florida in the United States are nesting earlier in response to warmer ocean
temperatures (Weishampel et al. 2004). Warmer waters also reduce the interval length between the
multiple clutches laid within a nesting season (Sato et al. 1998, Hays et al. 2002). Not all adult
turtles will breed each year, but the relative numbers arriving annually at widely separated rookeries
in Australia and the Indo-Pacific are similar, suggesting large-scale environmental forcing on
reproductive success (Limpus & Nicholls 1988, Chaloupka 2001). Variation in winter sea-surface
temperature anomalies partly explains internesting intervals of a Costa Rican population of green
turtles Chelonia mydas, with 2-yr remigration probabilities increasing in warmer years (Solow et al.
2002). In Australia, interannual fluctuations in numbers of green turtles nesting at rookeries within
the Great Barrier Reef are positively correlated with the Southern Oscillation Index, also with a
2-yr lag (Limpus & Nicholls 1988). Modelling studies suggest breeding intervals (time between
nesting years) are determined by resource provisioning on adult feeding grounds and the 2-yr lag
represents the time required for physiological provisioning for reproduction and migration (Hays
2000, Rivalan et al. 2005). Green turtles are herbivorous so are likely to be tightly coupled to
productivity in coastal waters (Broderick et al. 2001).
Mean egg-laying dates of many terrestrial bird species around the world have advanced con-
siderably in response to increasing temperatures (Archaux 2003, Both et al. 2004, 2005, Moller
et al. 2006). Migratory species are arriving earlier and leaving later (Mason 1995, Crick et al. 1997,
Lehikoinen et al. 2004, Marra et al. 2005, Jonzén et al. 2006). Most evidence is from the Northern
Hemisphere, but a similar pattern has recently been found in Australian migratory wetland birds
such as the curlew sandpiper Calidris ferruginea and the double-banded plover Charadrius bicinctus
(Beaumont et al. 2006). It is assumed that such changes are also occurring in Australian seabirds.
Protracted breeding seasons observed in seabird species in Western Australia are likely to be a
429
response to changing climate (Dunlop & Wooller 1986, Chambers et al. 2005). Breeding success
of little penguins Eudyptula minor in Bass Strait is correlated with sea temperatures and mean
laying dates are earlier in warmer years (Chambers 2004).
Pelagic systems
Physiology All plankton are poikilothermic and thus physiological rate processes and rates of
overall growth are highly sensitive to temperature (Eppley 1972, Peters 1983, Huntley & Lopez
1992), with many plankton having a Q10 between 2 and 3 (i.e., a doubling to tripling in the speed
of rate processes for a 10°C temperature rise). Species have a thermal optimum where growth is
maximal and thermal limits beyond which net growth ceases or becomes negative. Basal metabolic
losses increase with increasing temperature so that zooplankton fitness and, subsequently, abun-
dance and distribution may be adversely affected. Little information is available on temperature
ranges for Australian plankton, and in most cases experiments have been carried out with temperate
plankton strains. Culture studies do give some indication (e.g., Smayda 1976) and suggest that
species with tropical and subtropical distributions have growth optima <30°C. Optimal growth for
the dominant picophytoplankton species Synechococcus and Prochlorococcus in the Great Barrier
Reef is in the range 20–30°C (Furnas & Crosbie 1999), and in the Atlantic Ocean growth of
Synechococcus peaks at 28°C and growth of Prochlorococcus at a cooler temperature of 24°C
(Moore et al. 1995). As individual plankton strains have their own thermal optimum and limits for
growth, warming will have differential effects on the growth of individual species and changes in
phytoplankton and zooplankton community composition.
Although direct effects of temperature changes are fundamentally important to plankton rate
processes, indirect effects are also critical to plankton growth rates because zooplankton grow at
temperature-dependent maximal rates only when they are food saturated (Kleppel et al. 1996,
Hirst & Lampitt 1998, Richardson & Verheye 1998). Available evidence from tropical Australia
indicates that copepod growth and egg production rates are regulated primarily by food availability
rather than temperature (McKinnon & Thorrold 1993, McKinnon 1996, McKinnon & Ayukai 1996,
McKinnon et al. 2005). For example, generation times of the common coastal tropical copepod
Acrocalanus gibber decreased by 25% with a 5°C rise in temperature because of food limitation
(McKinnon 1996). Therefore, zooplankton growth rates appear to be severely food limited in the
warm, oligotrophic waters of tropical Australia (McKinnon & Duggan 2001, 2003). Climate impacts
on nutrient enrichment processes are thus likely to be at least as important in Australia as local
and direct temperature effects.
Temperature also has an effect on the body size of individual species of zooplankton. Copepod
body length typically decreases with increasing temperature (McKinnon 1996). Effects of temper-
ature on upper trophic levels may be strongly mediated by zooplankton size, which is a key
determinant of food quality for planktivorous fish. Warming of ocean waters will impact the
physiology or morphology of demersal and pelagic fish populations directly and indirectly, but too
little is known to speculate how these might be driven by climate change. Warming temperatures
will affect all life stages of these fish but egg and larval stages may be the most sensitive.
Distribution and abundance Plankton respond rapidly to ocean warming and have exhibited some
of the largest range shifts of any marine group (Hays et al. 2005). Members of the warm temperate
copepod communities in the northeast Atlantic have moved more than 1000 km poleward over the
last 50 yr (Beaugrand et al. 2002, Bonnet et al. 2005), although this may be more associated with
changing currents than warming. Concurrently, cooler water copepod assemblages have retracted
further toward the North Pole. It is likely that similar expansions have also occurred in warm
430
temperate and tropical dinoflagellates in the North Atlantic (M. Edwards 2004). Unfortunately,
plankton observations are rare in Australian waters. The only examples of plankton range extensions
are for the coccolithophorid Gephyrocapsa oceanica and the dinoflagellate Noctiluca scintillans.
Since the early 1990s this species has begun to appear in high densities off southeastern Australia,
with the likely cause being warmer sea temperatures (Blackburn & Cresswell 1993, Blackburn
2005, G. Hallegraef personal communication). Range expansions of other plankton species may
have considerable social and economic consequences. The box jellyfish Chironex fleckerii is cur-
rently at the southern limit of its range on North Queensland beaches where it causes problems for
bathers during summer; it may also expand its range further south as waters warm.
It is well recognised that sea temperature is a principal determinant of fish species abundance
and distribution (Lehodey et al. 1997, Roessig et al. 2004, Perry et al. 2005), biomass (Ware 1995,
O’Brien et al. 2000, Drinkwater 2005), and other critical life-history and physiological processes
(Burkett et al. 2001). Poleward shifts in distribution over the last century have been documented
for fish in the North Atlantic and the North Sea (Beare et al. 2004, Byrkjedal et al. 2004, Perry
et al. 2005, Rose 2005a,b), but observations from Australian waters are again few. Changes in the
distribution of large pelagic fishes, such as tunas and billfish, have been observed in response to
climate variability both seasonally (Zagaglia & Stech 2004) and interannually in terms of El Niño
Southern Oscillation (ENSO) (Lehodey 2001) and Rossby waves (White et al. 2004). Seasonal
distributions may be impacted if the timing of expansion or contraction of currents, such as the
Leeuwin or EAC, alters. For example, southern bluefin tuna Thunnus maccoyii are restricted to the
cooler waters south of the EAC and range further north when the current contracts up the New
South Wales coast (Majkowski et al. 1981). This response to climate variation has allowed real-
time spatial management to be used to restrict catches of southern bluefin tuna by non-quota holders
in the east coast fishery by restricting access to ocean regions believed to contain southern bluefin
tuna habitat (Hobday & Hartmann 2006). The seasonal presence of these fish along the east coast
of Australia may be reduced further if Tasman Sea warming continues. Preliminary analyses indicate
that changes may have already occurred, with fewer fish moving to the east coast in the Austral
winter (Polacheck et al. 2006).
Species from intermediate trophic levels (such as sardines and anchovies) are also crucial to
maintenance of biodiversity in the pelagic realm. These are particularly sensitive to climate impacts
based on studies elsewhere in the world (Chavez et al. 2003). A rare example from Australia is the
replacement in eastern Tasmania of cold-water jack mackerel Trachurus declivis with warm-water
redbait Emmelichthys nitidus from the EAC (Welsford & Lyle 2003), consistent with a warming
trend on the east coast of Australia and Tasmania.
Most species of marine turtles (except flatback turtles) move between coastal habitats and open
oceans, being distributed in waters generally warmer than 15–20°C (Davenport 1997), although
leatherbacks and loggerheads do penetrate into colder waters. Large leatherbacks are reported from
waters as cool as 8°C but juvenile leatherbacks (<100 cm carapace length) are rarely found in
waters <26°C (Eckert 2002). Reports from the Northern Hemisphere indicate that turtle populations
may already be responding to warmer temperatures. Most sightings of marine turtles in U.K. waters
over the past century are from the last 40 yr and sightings are increasing, suggesting a poleward
shift or expansion in distributions but may also be a result of better reporting (Robinson et al. 2005,
McMahon & Hays 2006). Global ranges of marine mammals are often related to water temperature
(Learmonth et al. 2006). However, climate-induced changes in prey availability will strongly influ-
ence distributions of marine mammals. A recent increase of warm-water cetaceans recorded in the
northeast Atlantic is likely to be the result of northward expansions linked to shifts of lower trophic
levels in response to warming temperatures (MacLeod et al. 2005).
431
Phenology There are insufficient data to assess changes in timing of plankton blooms in Australia,
but overseas studies show that timing is sensitive to climate warming and this can have effects that
resonate to higher trophic levels. In the plankton ecosystem of the North Sea, the timing of taxa
associated with low turbulent conditions in summer advanced with warming of 0.9°C from 1958
to 2002, with meroplankton moving forward by 27 days, dinoflagellates by 23 days, diatoms by
22 days, copepods by 10 days and non-copepod holozooplankton by 10 days (Edwards & Rich-
ardson 2004). These changes in phenology were greater than those observed in terrestrial commu-
nities (Root et al. 2003). Some groups such as dinoflagellates may not only be responding physi-
ologically to temperature, but may also react to temperature indirectly through earlier onset or
intensity of stratification. Others such as meroplankton are temperature sensitive because they are
dependent on temperature to stimulate physiological developments and larval release (Kirby et al.
2007). Important gelatinous meroplankton species that may display such tendencies include the
medusa stages of box jellyfish and the small highly poisonous Irukandji jellyfish, which has stings
that can be fatal to bathers. Only one species, Carukia barnesi, has been demonstrated to cause
Irukandji syndrome but at least six other, mostly undescribed, species may also be responsible in
Australian waters (Barnes 1964, Gershwin 2005, Little et al. 2006).
Although many plankton species are responding to climate warming, the magnitude of the
response differs throughout the community, having profound implications for the assembly, structure
and functioning of the pelagic communities and the entire pelagic ecosystem (Edwards & Rich-
ardson 2004). The different extent to which functional groups are moving forward in time in
response to warming (e.g., phytoplankton responding more than zooplankton) may lead to a
mismatch between successive trophic levels and a change in the synchrony of timing between
primary, secondary and tertiary production. Efficient transfer of marine primary and secondary
production to higher trophic levels such as commercially important fish species is largely dependent
on the temporal synchrony between successive trophic production peaks in temperate systems
(Cushing 1990). Thus, marine trophodynamics may have already been radically altered by ocean
warming and the extent to which this is happening in Australian temperate waters is unknown.
Phenology of migrations and spawning of many other marine species is also expected to alter. For
example, squid Loligo forbesi in the northeast Atlantic migrate to inshore spawning grounds earlier in
warmer years (Sims et al. 2001) while flounder Platichthys flesus migrate later (Sims et al. 2004).
Offshore benthic systems
Physiology Cold-water corals have been recorded from all the oceans and differ from shallow,
tropical, reef-forming species in that they lack symbiotic algae and are found at depths of several
hundred metres below sea level. Cold-water corals are restricted largely to temperatures between
4°C and 12°C (Roberts et al. 2003, Roberts et al. 2006). As these corals have evolved to be adapted
to this narrow yet stable temperature range, any rapid warming or cooling of temperatures is likely
to impact negatively on coral physiology. For example, rising temperatures will influence their
calcification rates, physiology and biochemistry.
Distribution and abundance Much of the relationship between temperature and benthic and dem-
ersal fish populations is likely to be a consequence of temperature-related productivity in pelagic
layers of the ocean, in addition to physiological dependencies. This relationship between temper-
ature and fish production and distribution is apparent over the decadal timescale where oceano-
graphic (temperature and productivity) regime shifts regulate zooplankton biomass, fisheries catches
and seabird abundances (Beamish et al. 1997, Mantua et al. 1997, McGowan et al. 1998, Beamish
et al. 1999, Koslow et al. 2002).
Range shifts of benthic and demersal fish species have already been observed in response to
warming of Australian waters. Most of these observations are from eastern or southeastern Australia,
432
although some such changes have been observed in Western Australia and it is not known whether
these differences are a reflection of differences in observation effort. Distributions of at least 36
species of Tasmanian marine fish have shifted poleward during the last decade (P. Last, personal
communication, CSIRO). Many of these are warm temperate reef species historically distributed
adjacent to the coast of New South Wales that have now become established south of Bass Strait.
Still others have shifted their ranges further south along the Tasmanian coast.
Phenology The benthic larval component of the zooplankton has shown large shifts in timing
compared with the holozooplankton in Northern Hemisphere temperate waters (Edwards & Rich-
ardson 2004, Greve et al. 2004). Evidence from the North Sea has shown that larvae of benthic
echinoderms are now appearing in the plankton about 6 wk earlier than 50 yr ago in response to
warmer temperatures. If Australian benthic systems responded similarly, peak larval abundances
of crown-of-thorns starfish could appear much earlier in the year, perhaps before the presence of
their normal predators (a potential positive feedback) or before wet season pulses in nutrients
originating from early wet season rains (a negative feedback).
Winds
Marine systems are influenced by wind fields, which drive major surface currents, and by episodic
wind events ranging in strength from low to extreme. In shallow waters, these wind events create
hydrodynamic disturbance whereas in deeper waters, wind fields and events contribute to hydro-
dynamic regimes that affect upwelling and hence productivity at different spatial and temporal
scales and across different trophic levels (Harris et al. 1991).
Coastal systems
Physiology Hydrodynamic stress will affect growth forms and morphological adaptations of plants
and animals (Denny & Gaylord 1996). For example, variation in the morphology of the kelp
Ecklonia radiata along the southern Australian coastline is related to wave exposure, longitude,
plant density and temperature at each site (Fowler-Walker et al. 2005, 2006). At sites with high
wave exposure, plants have longer stipes and smaller surface areas so are better adapted to cope
with high-energy water movement. Phenotypic responses to hydrodynamic stress are frequently a
trade-off between reducing mechanical damage and risk of dislodgement and obtaining nutrients/
food (Sebens 2002, Marchinko & Palmer 2003, Stewart & Carpenter 2003, Li & Denny 2004).
Populations cannot respond indefinitely to hydrodynamic stress so there are limits to the degree
of plasticity in morphological characteristics in response to the environment. Barnacles on Northern
Hemisphere exposed shores tend to have shorter cirri than those on sheltered shores (Arsenault
et al. 2001, Marchinko & Palmer 2003, Li & Denny 2004, Chan & Hung 2005) but above a threshold
current velocity barnacles cease to respond plastically to flow (Li & Denny 2004). Intertidal snails
tend to have thicker and/or larger shells on shores with high wave exposure (Frid & Fordham 1994,
Boulding et al. 1999). However, intertidal snails along the coast of southern Australia show no
differences in morphology with wave exposure, and it is hypothesised that the generally homoge-
neous and wave-exposed nature of Australia’s southern coastline may have favoured generalist traits
(Prowse & Pile 2005). Fauna and flora of Australia’s exposed southern coastline may be adapted
to cope with high variability in wave exposure.
Distribution and abundance Intertidal and shallow-water animal and plant communities are struc-
tured by wave exposure and local current velocity so species tolerant of high-energy hydrodynamic
forces dominate at high wave-exposed sites (Edgar et al. 1997, Coates 1998, Fonseca & Bell 1998,
Goldberg & Kendrick 2004, Fulton et al. 2005, Jonsson et al. 2006). An increase in wind strength
433
may increase wave exposure and may result in a considerable reduction in algal and seagrass
production or a shift in community composition in areas that are affected (Kendall et al. 2004,
Cruz-Palacios & van Tussenbroek 2005).
A general weakening of those winds is expected to hinder recruitment for coastal marine
populations. Strong relationships between wind strength and recruitment have been shown, includ-
ing in a coastal rocky reef fish (Heteroclinus sp.) for which enhanced settlement followed wind-
driven productivity boosts (Thresher et al. 1989). Prolonged periods of strong winds have impacted
the breeding success of the sooty tern Sterna fuscata and common noddy Anous stolidus in the
Great Barrier Reef region with large-scale desertion of nests and starvation of chicks (King et al.
1992). Environmental conditions associated with strong winds may have led to a reduction in prey
availability or a reduction in the foraging success of adults. Nests were also lost through inundation
by waves and shoreline erosion (King et al. 1992).
Phenology Winds and waves have the potential to affect timing of the reproduction of algae. For
example species of Fucus in the North Atlantic release spores only under calm conditions at low
tide at certain times of the year (Brawley 1992, Serrão et al. 1996, Brawley et al. 1999). It is not
clear whether this is an absolute condition for reproduction, or whether it is simply periods of
relative calm that are required. It is also unknown whether any Australian species of marine plants
have similar requirements for reproduction. It has also been suggested that the timing of mass
spawning in tropical reef corals is related to seasonal wind and current fields, coinciding with times
of the year when calm conditions are likely to occur (Babcock et al. 1994). It is thought that the
fertilisation success of coral populations may be the ultimate factor responsible for this pattern
(Oliver & Babcock 1992), so any change in the seasonal wind pattern may affect reproduction and
recruitment. If climate change decouples factors such as seasonal wind patterns and seasonal
temperature cues that may be important for mass spawning corals then the reproductive success of
these species may be reduced.
Pelagic systems
Distribution and abundance Wind is one of the driving forces of currents and vertical mixing in
the water column. Wind therefore affects mixing depth and intensity and may thus be seen as a
proxy for mixing depth, mixing intensity, and light and nutrient supply to the surface layer. Climate
models consistently project a poleward shift in the zonal winds that normally cross the southern
part of Australia, and these projections are consistent with recent changes in the Antarctic Oscillation
Index (Gillett & Thompson 2003; also see Jones & Widmann 2004). The projected general weak-
ening of those winds following this shift may reduce recruitment to marine fish populations. Strong
relationships between wind strength and recruitment exist for some species, such as the commer-
cially exploited blue grenadier Macruronus novaezelandiae in outer continental shelf waters
(Thresher et al. 1992). In southeastern Australia, Harris et al. (1992) found evidence that reduced
production of the jack mackerel Trachurus declivis off Tasmania resulted from decreased wind
stress and subsequent decreases in large zooplankton.
Distribution and abundance The variability in the annual frequency of strong zonal westerly
winds has been related to catch rates and recruitment variability in several southeastern demersal
fisheries (Harris et al. 1988). The collapse of the gemfish fishery Rexea solandri in that region was
likely a consequence of the combination of weak recruitment due to declining winds and overfishing
(Thresher et al. 1996). A variety of southeastern shelf teleosts exhibit a decadal-scale recruitment
cycle, in several cases directly linked to regional wind fields (Thresher 2002, Jenkins 2005).
434
Ocean currents
Currents and ocean circulation systems strongly affect dispersal, migration and geographic distri-
bution of species and therefore have implications for the connectivity of marine systems. Southward-
moving currents such as the EAC and Leeuwin Current interact with southern coastal and offshore
waters, influencing temperature and regional productivity (Harris et al. 1987, Ridgeway & Dunn
2003, Ridgway & Condie 2004).
Coastal systems
Distribution and abundance Many marine plants and animals rely on water movement for dis-
persal, particularly for early life stages. Distributional patterns of marine populations often reflect
connectivity of marine systems. Evidence is mounting that despite the potential for long-distance
dispersal, actual dispersal distances for coastal fauna may be constrained by behavioural mecha-
nisms such as vertical migration. Typical larval dispersal distances for coral reef fish in the
Caribbean are on a scale of 10–100 km, with dispersal distances strongly determined by active
movement of larvae (Cowen et al. 2006). Some coastal invertebrates have very short larval durations
which will restrict dispersal distance (McShane et al. 1988, Sammarco & Andrews 1988, Davis &
Butler 1989, Stoner 1992). Further, the viability of larvae and plant propagules may diminish over
time. For example, propagules of the mangrove Avicennia marina may only be able to establish
successfully within the first 4–5 days of dropping (de Lange & de Lange 1994). The southern limit
of this species in New Zealand appears to be controlled by limited transport by coastal drift and
lack of suitable habitat within the dispersal range of existing populations, rather than by climatic
factors (de Lange & de Lange 1994).
The most southerly mangroves globally are found at Corner Inlet in Victoria (de Lange & de
Lange 1994). These may be relict populations from when favourable climate extended further south
than at present. Projected global warming and strengthening of the EAC may facilitate further
southerly expansion of mangrove species. Alternatively, this southerly limit may be set (and
restricted) by eastward water movement through the Bass Strait (de Lange & de Lange 1994).
Southward water movement through the Bass Strait by wind-induced drift is slow and is insufficient
to transport the propagules to Tasmania within the 5-day period for viable establishment (de Lange &
de Lange 1994, Clarke et al. 2001). Therefore, even if temperatures in Tasmania become warm
enough to support Avicennia populations (conventional wisdom is that latitudinal range edges of
mangroves are determined mainly by freezing temperatures) they are unlikely to become established
there. Currents thus act as a barrier as well as an aid to dispersal for many marine organisms and
therefore determine adult abundances as well as distributional limits (Gaylord & Gaines 2000).
Incidentally, this means that a key consequence of future climate change will be influences on
current patterns, often on a small scale and therefore dependent on fine-scale variations in weather
and current patterns, which are still difficult to predict.
In addition to affecting the range of species distributions, a change in the strength of currents
may alter the overall strength of recruitment. One of the best examples of this comes from the
correlation between the strength of the Leeuwin Current and recruitment of the western rock lobster
Panulirus cygnus. The strength of the Leeuwin Current is highly correlated with ENSO and in
El Niño years when the current is weak, rock lobster recruitment is also weak (Caputi et al. 2001).
The mechanism underlying this is not well understood, but it is clearly more complex than simply
a range extension and may be related to temperature or cross-shelf transport, mixing and productivity
driven by the Leeuwin (Griffin et al. 2001).
The establishment of long-spined sea urchins Centrostephanus rodgersii in Tasmania in the
1960s has been attributed to larval transport from northern populations by the EAC (Johnson et al.
2005). Populations have since expanded in Tasmania and have resulted in the elimination of
435
macroalgae in some areas through intense grazing pressure. A reduction in the density of rock
lobster Janus edwardsii and abalone Haliotis rubra in areas devoid of macroalgae has serious
implications for fisheries targeting these species. It must be assumed that any major shift or
strengthening of wind fields and major currents may have profound implications for Australian
coastal organisms.
Pelagic systems
Distribution and abundance Transport by large ocean currents plays a major role in the move-
ments of marine turtle hatchlings and early juveniles to ocean pelagic nursery habitats where young
turtles remain for a number of years exploiting biologically rich environments linked to current
systems and convergence zones (Carr 1987, Witherington 2002, Ferraroli et al. 2004). Juvenile and
adult turtles undertake extensive migrations; juvenile loggerheads originating from Australian
populations have been identified from feeding grounds off Baja California, representing a journey
that crosses the entire Pacific Ocean aided by the North Pacific Current (Bowen et al. 1995). Adult
loggerheads and leatherbacks forage at fronts and eddies and are associated with major currents
(Ferraroli et al. 2004, Polovina et al. 2004). Turtles have also been tracked swimming against
prevailing currents as well as with currents so may only use current flows opportunistically to
facilitate transport (Luschi et al. 2003, Polovina et al. 2004). Alteration of major current systems
will impact the navigational abilities of marine turtles and deflect turtle movements (Luschi et al.
2003, Robinson et al. 2005).
Distribution and abundance Cold-water corals are found in areas of fast currents and this is
evident on Australian seamounts where corals occur in distinct depth zones (Koslow et al. 2001).
Alteration of currents may make areas unfavourable for coral growth and, given the low growth
rate, colonisation of newly available areas with optimal environmental conditions may be slow and
may take many decades before a viable population size is reached. Fast flow may also be necessary
for larval supply or retention to establish or maintain populations (Genin et al. 1986). Changes in
local current regimes could alter the ‘stepping stone’ function of seamount chains, whereby the
biology on distant seamounts is linked by intermediate ones, and have a considerable impact on
coral distribution (Roberts et al. 2003). Survival of cold-water corals appears to be controlled by
oceanographic conditions. Chemical analysis of deep-water corals off southern Australia has indi-
cated a long-term deep cooling that commenced in the mid-eighteenth century and is a result of
enhanced poleward flow of the warm EAC as it interacts with the colder subsurface countercurrents
(Thresher et al. 2004). This strengthening of the EAC is predicted to continue as the global climate
warms (Cai et al. 2005, Cai 2006) with considerable impacts for ocean circulation and marine
biodiversity, including cold-water coral ecosystems.
Mixed-layer depth and stratification

Mixing depth and mixing intensity in the surface ocean and the associated stratification are key
factors for the production of phytoplankton and of higher trophic levels because they fundamentally
affect the supply of nutrients (from below) and light (from above), and sinking losses of phyto-
plankton (Mitchell & Holm-Hansen 1991, Huisman & Weissing 1995, Diehl 2002) and because
consumer biomass is positively related to the productivity of their food (Grover 1997).
436
Pelagic systems
Distribution and abundance Experimental manipulation of mixing depth has demonstrated the
positive impact of decreasing mixing depth (increasing light supply) on the biomass of temperate
phyto- and mesozooplankton when nutrients are relatively abundant (Kunz 2005). Flagellates
profited from shallower mixing and the heterotroph-to-autotroph (mesozooplankton-to-phytoplank-
ton) biomass ratio was higher at low mixing depth. Variability of mixing depth and change in
stratification in several ocean regions since the 1950s provide striking examples of potential impacts
on pelagic communities. In the central and subarctic North Pacific Ocean, large variability in
plankton primary and secondary production has been linked to a decadal-scale climate change event
between the mid-1970s and the late 1980s and associated changes in the depth of the winter and
spring mixed layers (Venrick et al. 1987, Polovina et al. 1995, Hayward 1997). These impacts on
lower trophic levels appear to have propagated to higher trophic levels with pelagic larvae, including
squid, salmon and flying fishes with different mechanisms operating in different regions (Polovina
et al. 1994, 1995). In the northwest Hawaiian Islands (situated in the North Pacific subtropical
gyre), chlorophyll concentration and primary production were positively related to deepening of
the mixed layer due to increased nutrient supply (Venrick et al. 1987, Polovina et al. 1995) while
in the subarctic Gulf of Alaska copepod abundance and, likely, primary production were positively
related to shallowing of the mixed layer due to increased light availability (Polovina et al. 1995).
In the Northern California current, a decrease in macrozooplankton biomass by 80% since 1951
has been related to reduced nutrient transport across the thermocline due to warmer sea-surface
temperatures and increased stratification (Roemmich & McGowan 1995). In the North Atlantic,
large-scale northward shifts in the distribution of warm-water phyto- and zooplankton and changes
in the abundance of plankton between 1958 and 2002 have been related to increasing water column
stratification (Richardson & Schoeman 2004, Hays et al. 2005, Edwards et al. 2006).
With the projected enhancement of stratification around most of continental Australia nutrient
transport to the surface layer will be reduced over vast areas of the pelagic zone. Most Australian
waters are therefore likely to become more depauperate in nutrients with repercussions for produc-
tion and biomass of most pelagic (and benthic) food webs. Cyanobacteria, flagellates and dinoflagel-
lates (including nuisance and harmful algal bloom species) may increase in abundance where
vertical mixing decreases and the ‘microbial loop’ may be favoured over the relatively more
productive ‘classic’ food web in affected areas. The productive temperate pelagic province may
shrink considerably in area and potentially become restricted to west of Tasmania by 2100. In
tropical surface waters where increasing stratification lifts the oxycline, the abundance of pelagic
apex predators, such as skipjack and yellowfin tuna may decline.
Phenology In the North Atlantic, earlier timing of dinoflagellate blooms in spring is partly
attributed to earlier and enhanced stratification (Edwards & Richardson 2004, Richardson & Schoe-
man 2004). In Tasmanian waters, zonal westerly winds stimulate deeper and/or stronger vertical
mixing and affect the timing and duration of phytoplankton blooms (Harris et al. 1988).
Distribution and abundance Deep seafloor habitats, with the exception of hydrothermal vents and
cold seeps, are typically areas of low productivity, relying on the flux of detritus from surface
waters which is partially regulated by mixed layer depth. Despite this, species diversity can locally
be very high (Snelgrove & Smith 2002). Seamounts, with topographically enhanced currents are
areas of high productivity (e.g., Koslow 1997) but are still sensitive to the flux of organic matter
from surface waters, albeit over a wider area than that of the seamount itself. This coupling to
437
surface productivity may mean the deep sea is particularly susceptible to climate change (Glover
& Smith 2003). If surface productivity is reduced as climate warms then the reduction in organic
carbon flux to the sea floor will lead to a reduction in benthic biomass.
CO2, pH and calcium carbonate saturation state

Changes to the atmospheric concentration of CO2 and hence carbonate ions represents a serious
threat to calcifying organisms such as corals, pteropods and coccolithophores (Raven et al. 2005),
especially as calcification of most organisms appears linearly related to the carbonate ion concen-
tration (Langdon et al. 2000). The level of calcification is significant in that it represents concen-
trations at which organisms such as tropical reef-building corals no longer calcify. Marine organisms
differ in their susceptibility to acidification depending on whether the crystalline form of their
calcium carbonate is calcite (calcifying phytoplankton, foraminiferans) or aragonite (pteropods,
corals). Calcite is less soluble than aragonite, making it less susceptible to pH changes. Other
effects of increased CO2 on the physiology of marine flora and fauna are less well understood.
Experiments to determine the likely response of marine organisms to pH changes have explored
large changes in pH (>1.0) under laboratory conditions (Kikkawa et al. 2003, Pedersen & Hansen
2003a,b, Pörtner et al. 2004, Engel et al. 2005) but little is known on what the gradual long-term
effects of pH lowering will be on marine organisms.
Coastal systems
Physiology Marine plants vary in their degree of immersion in water, from mangroves that
generally have their foliage and flowers above the water to macroalgae and seagrasses that are
either fully submerged or submerged for part of the tidal cycle. Land plants, including mangroves,
capture CO2 primarily by diffusion so that increasing atmospheric CO2 generally hastens photo-
synthesis, productivity and growth (Ainsworth & Long 2005). Seagrasses, although submerged,
are of terrestrial origin and so rely primarily on dissolved CO2; thus they are photosynthetically
inefficient in sea water (Invers et al. 1997, 2002, Short & Neckles 1999). By contrast, most marine
phytoplankton and macroalgae have mechanisms that actively concentrate and take up inorganic
carbon as CO2, bicarbonate ions (HCO3−) or both, so changes in dissolved CO2 have less effect on
their rates of photosynthesis (Giordano et al. 2005). Carbon-concentrating mechanisms are not as
common in benthic photosynthetic organisms (Giordano et al. 2005).
Mangrove growth may be stimulated as CO2 levels increase. Seedlings of Rhizophora mangle
grown under double ambient CO2 for a year exhibited increases in growth and photosynthetic rate
(Farnsworth et al. 1996). The young plants also became reproductive a year earlier than in the field,
so elevated CO2 may accelerate maturation as well as growth (Farnsworth et al. 1996). However,
the long-term response of mature mangrove forests to elevated CO2 is unknown. A widespread
thickening of terrestrial vegetation observed in parts of Australia may be induced by recent climate
change although is more likely the result of changes in land use (Bowman et al. 2001, Australian
Greenhouse Office 2003).
Australian coastal waters are generally low in phosphate and nitrate but as seagrasses are rooted,
they can take up these essential nutrients from the sediment. Therefore, seagrasses are primarily
carbon limited. An increase in atmospheric CO2 will result in a higher proportion of dissolved CO2
in the oceans, potentially increasing seagrass biomass, deepening of seagrass depth limits and
enhancing of the role of seagrass beds in carbon and nutrient cycles (Zimmerman et al. 1997, Invers
et al. 2002). Intertidal macroalgae, which generally use bicarbonate when submerged, may only
benefit from elevated CO2 during aerial exposure (Farnsworth et al. 1996, Beardall et al. 1998, Gao
et al. 1999, Zou & Gao 2002, 2005).
438
Coral reefs represent a balance between calcification and erosion, with 90% of what is laid
down by calcifiers being removed by erosion. Ocean acidification could tip the balance from net
calcification to erosion. If atmospheric CO2 levels reach 500 ppm, projected to occur by the end
of this century, then coral viability will be severely compromised (Hoegh-Guldberg 2004). At low
carbonate ion concentrations (<200 µmol kg−1), calcification of corals and many other calcifying
organisms effectively becomes zero. The actual seriousness and time frame of these changes have
yet to be properly assessed.
There has been some debate about the significance of the threat of ocean acidification to the
long-term viability of coral reefs (see McNeil et al. 2004 vs. Kleypas et al. 2005). Changes in
calcification rates over recent centuries estimated from cores from long-lived corals such as Porites
on the Great Barrier Reef show evidence of an increase in calcification rates over the 50 yr prior
to 1982 (Lough & Barnes 2000). Calcification rates were highly correlated with average sea
temperature, with an annual average increase in calcification of 0.3 g cm−2 yr−1 for each degree of
ocean warming. Lough and Barnes (2000) suggested the increase in calcification was probably due
to the 0.25°C warming of sea temperature on the Great Barrier Reef over the last 50 yr. Although
calcification does increase with temperature, it does not increase indefinitely; several studies have
shown that it increases up to the summer sea-temperature maximum, but declines rapidly at warmer
temperatures. Interactions between temperature and decreasing pH are still largely unknown but
are likely to be considerable given, for example, the linkages between metabolic rate (which is
temperature sensitive) and calcification. Most authors have concluded that the combination of the
two pressures on calcifying organisms such as corals will be largely negative and synergistic
(Hoegh-Guldberg 2004).
Acidification may be expected to increase physiological stress on other calcifiers. Metabolic
efficiency and growth rates of bivalves and other molluscs will be impaired (Michaelidis et al. 2005,
Berge et al. 2006). Experiments have also shown that under lowered pH conditions the fertilisation
rate of eggs of intertidal echinoderms declined and larvae were severely malformed (Kurihara et al.
2004).
Distribution and abundance Macroalgae take up primarily bicarbonate ions for photosynthesis.
As only a small proportional change in bicarbonate concentration will occur as atmospheric CO2
levels rise, little enhancement of growth is expected (Beardall et al. 1998). However, increased
acidification of the oceans may have severe consequences for coralline algae (Gao et al. 1993)
therefore enhancing competitive advantages of non-calcifying species over calcifying species (Gao
et al. 1993, Beardall et al. 1998).
Potential range shifts in tropical corals with warming may be restricted by the future latitudinal
gradient in the carbonate saturation state of sea water (Figure 6). The undersaturation of aragonite
and calcite in sea water is likely to be more acute and happen earlier further south in the Southern
Hemisphere and then move northward (Orr et al. 2005). This means that corals will not be able to
move further south into cooler waters in response to warming seas because these waters are likely
to be undersaturated in calcium carbonate. The additional problem of reduced light levels at higher
latitudes is also probably an important limit in this respect.
Pelagic systems
Physiology Seawater pH affects phytoplankton via several processes. pH is an important deter-

minant of the growth rate of phytoplankton species, with some growing consistently well over a
wide range of pH and the growth rate of others varying considerably over a pH range of 7.5–8.5
(Hinga 2002). Both temperate and tropical coccolithophorids show reduced calcite production and
an increased proportion of malformed liths at decreased pH (Riebesell et al. 2000, Engel et al. 2005).
439
Declining pH may also alter the growth rates of photosynthetic organisms; in particular changes
in pH will affect the kinetics of the uptake of nutrients. Nitrification in marine bacteria is negatively
affected below a pH of ~8. Because nitrification is an important pathway of nitrate supply to
phytoplankton, nitrate availability for phytoplankton is likely to be reduced at pH <8.0, with
consequences for phytoplankton community composition and productivity (Huesemann et al. 2002).
Decreasing pH has also been found to increase the availability of potentially toxic trace elements
such as copper, which may affect phytoplankton survival (Kester 1986). Changes may also occur
in cell composition, which could affect the nutritional value of the microorganisms to the animals
that feed on them.
As phytoplankton have carbon-concentrating mechanisms, photosynthesis is generally not
carbon limited, even at present CO2 levels. In almost all phytoplankton species, doubling CO2
concentration only increases photosynthesis by <10% (Beardall & Raven 2004, Schippers et al.
2004, Giordano et al. 2005). The small number of studies that have investigated effects of CO2 on
phytoplankton community composition suggested that elevated CO2 concentrations favour diatoms
over flagellates and coccolithophores (Antia et al. 2001, Tortell et al. 2002).
The physiology of larger animals such as fish and squid are likely to be influenced by increasing
CO2 levels in the oceans, which influences tissue acid-base regulation and thus metabolism. Squid
are acutely sensitive to even small changes in ambient CO2 due to their high metabolic oxygen
demand for locomotion (jet propulsion) and a strong relationship between O2 binding in the blood
and pH (see Pörtner et al. 2004 ). Pelagic fish generally have lower metabolic rates and some venous
oxygen reserve so are only moderately sensitive to changes in ambient CO2. The projected increases
in CO2 are below lethal threshold levels; synergistic effects of warmer temperatures and increased
CO2 may influence growth and reproduction in large pelagic fauna.
Distribution and abundance The direct effect of ocean acidification on calcifying zooplankton
will be to increase shell maintenance costs and reduce growth. Pteropods with their aragonite shells
are particularly vulnerable to ocean acidification (Orr et al. 2005). In the Southern Ocean, shelled
pteropods are prominent components of the food web contributing to the diet of carnivorous
zooplankton, myctophid and other fishes, and baleen whales, as well as forming the entire diet of
gymnosome molluscs. Pteropods can also account for the majority of the annual export flux of
both carbonate and organic carbon in the Southern Ocean. Shells from live pteropods dissolve rapidly
when placed in water undersaturated with aragonite, similar to the levels that are likely to exist in
2100. If pteropods cannot grow their protective shell, then their populations are likely to decline
and their range will contract toward lower-latitude surface waters that remain supersaturated in
aragonite. In Australian waters, pteropods are relatively rare but can be locally common. For
example, the pteropod Cavolinia longirostris can form dense aggregations on the Great Barrier
Reef during summer (Russell & Colman 1935), occurring in such large numbers that their shells
wash up on beaches (D. McKinnon, personal communication).
Distribution and abundance Decreases in ocean pH will directly impact offshore soft sediment
organisms that rely on calcium carbonate structures such as molluscs and foraminiferans. Changes
in pH will also impact benthic organisms by influencing the composition of sediment as a large
fraction in Australia is calcium carbonate in origin. For example, foraminiferan remains constitute
most of the sediments in sandy regions of the Great Barrier Reef and a decline in the abundances
of pelagic and benthic foraminiferans is likely to reduce the sedimentation of their skeletons to the
sea bottom (McKinnon et al. in press).
The global distribution of cold-water corals is influenced by seawater carbonate chemistry with
a clear relationship between the occurrence of cold-water scleractinian corals and depth of the
440
aragonite saturation horizon (Guinotte et al. 2006). The aragonite saturation horizon represents the
limit between the upper saturated and the deeper undersaturated waters; calcium carbonate can
form above the horizon but dissolves below it (Raven et al. 2005). As atmospheric CO2 levels
increase, the depth of the aragonite saturation horizon will rise closer to the ocean surface and the
entire Southern Ocean water column could become undersaturated by 2100 (Caldeira & Wickett
2005, Orr et al. 2005). Cold-water corals are thus likely to be much more vulnerable to changes
in ocean chemistry than shallow tropical reef-building corals (Raven et al. 2005). Over the next
100 years, the predicted decrease in the aragonite saturation horizon in the oceans will result in
only 30% of known deep-sea coral reefs and mounds remaining in supersaturated waters compared
to the present-day figure of >95% (Guinotte et al. 2006). Ocean waters south of Australia may become
inhospitable for cold-water corals below a few hundred metres. The shallowest pinnacles in the
Tasmanian Seamounts Marine Reserve peak at about 600 m below the surface so cold-water corals
on these seamounts could simply disappear, along with their multitude of associated organisms.
Solar radiation
Future changes in UVR are difficult to predict (see p. 416) but in the following sections we discuss
the potential impacts of heightened UVR, while recognising that the possibility of ozone recovery
is becoming more likely. The extent to which UVR affects marine organisms will depend on factors
such as aerosol concentrations, cloud cover and the concentration of dissolved and particulate matter
in the water column (Jerlov & Steeman-Nielsen 1974, Smith & Baker 1979, Erga et al. 2005).
Coastal systems
Physiology UVR damages DNA and causes photo-oxidative stress in plants and animals (Setlow
& Setlow 1962). In the aquatic environment, UVR effects should be most intense near the water
surface. Sessile species, such as intertidal fauna and corals, do not have the capacity to avoid UVR
through evasive movements and so can be exposed to powerful solar irradiances, particularly in
tropical waters (see Shick et al. 1996). Tropical reef corals with their symbiotic zooxanthellae need
to be exposed to sunlight for photosynthesis, so they are adapted for a high-UVR environment.
However, an increase in UV light may exacerbate the simultaneous stress of warmer temperatures
on corals and thus contribute to coral bleaching (Lesser 1996, 1997, Baruch et al. 2005, Drohan
et al. 2005). Sublethal effects of UVR include depressed calcification and skeletal growth in corals
(see Shick et al. 1996).
Increased UVR reduces plant photosynthetic efficiency and biomass (Dawson & Dennison
1996, El-Sayed et al. 1996, Moorthy & Kathiresan 1997, 1998). Excessive UVR can induce
photoinhibition in dinoflagellates, macroalgae, seagrasses, and the symbiotic zooxanthellae of
tropical corals and sea anemones, with tolerances varying among species and life stages (Dawson &
Dennison 1996, Graham 1996, Bischof et al. 1998, Häder et al. 1998; also see Shick et al. 1996).
For example, net photosynthesis of the mangrove Rhizophora apiculata seedlings increased by
45% for a 10% increase in UVR but a 59% decrease in net photosynthesis occurred with a 40%
increase in UVR (Moorthy & Kathiresan 1997). Many tropical species may already be near their
upper limits of UVR tolerance so any further increase may reduce the ability of vulnerable species
to persist near the water surface, thus leading to a shift in community composition (Häder et al.
1998).
Egg and larval stages of many marine invertebrates and fish are highly susceptible to UV
damage, particularly those that are pelagic (Lesser et al. 2003, Wellington & Fitt 2003, Przeslawski
et al. 2004, 2005, Bonaventura et al. 2006). Increased UVR is known to have a deleterious effect
on some adult fish, damaging ocular components and the epidermis, depressing the immune system,
441
and allowing invasion of pathogens (Zagarese & Williamson 2001, Markkula et al. 2005). Some
coral reef fishes that are exposed to intense irradiance are able to sequester UV-absorbing com-
pounds from prey and thus are less vulnerable to UV increases (Zamzow 2004).
Distribution and abundance Intertidal and subtidal algae and seagrasses will generally be sus-
ceptible to changes in solar irradiance. Upper depth limits of many species in these groups may
deepen or grow shallower with increased or decreased levels of UVR, respectively (Dawson &
Dennison 1996). Early life stages may be more susceptible to UVR than mature plants, thus
regulating depth limits of adults (Graham 1996, Rijstenbil et al. 2000, Swanson & Druehl 2000,
Cordi et al. 2001). For example, the upper depth limit of some kelp species is determined by
susceptibility of their zoospores to UVR (Swanson & Druehl 2000, Wiencke et al. 2006) or early
post-settlement stages (gametophytes or embryonic sporophytes) to photosynthetically active radi-
ation (Graham 1996). Plants produce UV-absorbing compounds found predominantly in the epi-
dermis and there is some capacity for adaptation in certain species. Levels of UVR-blocking pigment
in certain tropical seagrasses increase when plants are grown at higher irradiance (Abal et al. 1994,
Detres et al. 2001).
Unlike sessile plants and animals, mobile fauna can shift distributions or retreat to refugia
during periods of high solar radiation. One example is that the settlement of coral larvae is influenced
by UVR levels (Kuffner 2001, Gleason et al. 2006), so these larvae may have some choice in
settlement locations. How alteration of solar radiation patterns will affect behavioural responses of
other marine animals is, however, generally difficult to predict. Visual systems of some shallow-
water fishes use UV wavelengths and allow con-specific communication during breeding, shoaling
or territorial behaviour (Losey et al. 1999, 2003, Garcia & de Perera 2002, Losey 2003, Siebeck
2004, Modarressie et al. 2006). It is assumed there is large plasticity in behavioural responses so
at least some populations may adapt rapidly.
Interspecific variability in the capacity of marine plants and animals to adapt to UVR changes
(Hanelt et al. 1997, Choo et al. 2005) may lead to shifts in shallow-water and coral reef community
structure if solar irradiance changes. Effects on communities should be most pronounced where
there is a strong differential sensitivity to UVR between species or where protection against UVR
is metabolically expensive or juvenile stages are found near the water surface (Wahl et al. 2004).
Pelagic systems
Physiology In phytoplankton, UVR can negatively impact several physiological processes and
cellular structures, including photosynthesis, carbon and nutrient uptake, the ratio of polyunsatu-
rated to saturated fatty acids, cell motility and orientation, the DNA, and life-span (Behrenfeld
et al. 1993, Goes et al. 1994, Hessen et al. 1997, Wilhelm et al. 1997, Garde & Cailliau 2000,
Hogue et al. 2005, Litchman & Neale 2005). These effects not only reduce phytoplankton growth,
production and biomass (Worrest et al. 1978, Döhler 1994, Hessen et al. 1997, Keller et al. 1997,
Wängberg et al. 1999) but also compromise the ability of phytoplankton to adapt to changing
environmental conditions and respond to possibly hazardous situations (Häder & Häder 1989, Häder
& Liu 1990). Although some phytoplankton are capable of acclimating to UVR via increased
pigmentation or capability to repair damaged DNA, this inevitably involves metabolic costs reducing
the energy that would otherwise be available for cell growth and division (Häder et al. 1998, Garde
& Cailliau 2000). Increases in the cellular carbon-to-nutrient ratio and cell size reduce the nutritional
value of phytoplankton for grazers. Negative effects of altered food quality are known to propagate
to higher trophic levels and have been related to reductions in the abundance of copepod nauplii
in experimental mesocosms (Hessen et al. 1997, Keller et al. 1997).
UVR may positively affect bacteria and phytoplankton production because it increases the
photolysis of dissolved organic carbon and colloids thereby increasing the availability of essential
442
plant macro- and micronutrients from such compounds (Rich & Morel 1990, Palenik et al. 1991,
Wängberg et al. 1999).
There have been relatively few studies on the effects of UVR on marine zooplankton, in
comparison with studies on phytoplankton. UVR is also known to damage various life stages of
zooplankton such as copepods and shrimp, as well as eggs and larvae of crabs and fish that are
temporary members of the plankton (Hunter et al. 1982, Damkaer & Dey 1983, El-Sayed et al.
1996, Kouwenberg et al. 1999a,b). In copepods, UVR has been found to lower fecundity, increase
mortality and affect the sex ratio (Karanas et al. 1979, 1981, Bollens & Frost 1990). Because UVR
can have deleterious consequences for those organisms that lack photoprotective mechanisms, many
of the permanent members of the neustonic copepod community which are common in Australia’s
warmer waters have pigments to reduce such damage. The effects of UVR radiation on fish eggs
or larvae have rarely been investigated (see Zagarese & Williamson 2001). The few existing studies
found deleterious effects of UVR in clear waters and confirm the importance of dissolved organic
carbon in ameliorating those effects (Hunter et al. 1982, Keller et al. 1997, Zagarese & Williamson
2001).
Distribution and abundance The degree of water column stratification crucially affects the expo-
sure to UVR of those plankton that do not migrate vertically. Shallowing of the mixed surface layer
and more energetic turbulence both increase plankton exposure to UVR and, therefore, their chance
to receive harmful doses (Keller et al. 1997, Garde & Cailliau 2000, Barbieri et al. 2002, Hernando
& Ferreyra 2005). Differential sensitivities of individual plankton taxa are thus likely to cause or
have caused shifts in community structure or even ecosystem integrity, depending on the magnitude
of changes in UVR. It is questionable to what extent the observed short-term effects of UVR on
individual organisms can be used to estimate long-term ecosystem response (Häder et al. 1998).
Precipitation and storms

A 70-yr drying trend has occurred along the eastern seaboard of the Australian mainland (Australian
Greenhouse Office 2003), and this has reduced vegetation and destabilised sediment in coastal
watersheds. Following intense storms in temperate Australia and monsoonal rains in tropical
Australia, runoff from rivers tends to carry high sediment loads. The increasing frequency of intense
storms as a result of climate change (Emanuel 2005, Webster et al. 2005) is likely to increase extreme
rainfall events, hence altering runoff of freshwater and suspended sediment loads. There are clearly
numerous indirect impacts associated with changes in freshwater flux in addition to the direct
potential for fresh water to have physiological effects on marine organisms.
Coastal systems
Physiology Changes in rainfall patterns, and associated changes in watershed geomorphological

dynamics, will affect the dynamics of coastal marine ecosystems through the physiological effects of
large-scale flooding, fluctuations in salinity, and increases in turbidity and nutrients on resident
organisms such as mangroves and wetland flora. Mangroves are adapted for coastal areas with
waterlogged and often anoxic soils but their tolerance of salinity stress varies among species. As
salinity levels increase, mangroves are faced with increasing salt levels in the tissues (see Field 1995).
A ‘zonation’ of mangrove species can generally be observed going from the sea to the land or upriver
from the mouth of estuaries reflecting the ecophysiological response of the plants along these and
other environmental gradients and disturbance regimes (Ellison & Farnsworth 1993). Rainfall directly
influences the salinity of the intertidal waters and sediments but also influences salinity through
freshwater runoff from the land and freshwater seepage into the soil (Twilley & Chen 1998).
443
Hydrology of mangroves is complex; tidal inundation, rainfall, groundwater seepage and evap-
oration all influence soil salinity and have a profound effect on mangrove growth. Hydrology model
simulations of mangrove systems in southwest Florida have demonstrated that mangroves in the
upper intertidal are particularly sensitive to reductions in rainfall, even though these are areas with
minor freshwater input (Twilley & Chen 1998). Animals and plants living in the upper intertidal
are generally near the upper limits of environmental tolerance limits so small alterations in climate
may have a greater impact on upper shore organisms than in the lower intertidal. For example,
seedlings of the mangrove Rhizophora apiculata grew more rapidly in the lower intertidal than
those in the upper intertidal, presumably reflecting the additional stresses in the upper intertidal
(Kathiresan et al. 1996).
Freshwater runoff can increase sediment loading of coastal waters thus imposing metabolic
costs on corals and other organisms that can potentially reduce growth or lead to mortality in severe
cases (see Fabricius 2005). Other effects of sediment on corals can occur at early life-history stages.
Sediment has been shown to reduce coral fertilisation (Gilmour 1999), as well as settlement
(Babcock and Davies 1991) and post-settlement survival of recruits (Babcock and Smith 2002).
Pollutants that are carried with flood waters are also known to have detrimental effects on the early
life-history stages of corals. For example the herbicide diuron, commonly used in catchments
adjacent to the Great Barrier Reef, inhibits coral metamorphosis and settlement (Negri et al. 2005).
Distribution and abundance Mangroves are considered highly susceptible to alteration in rainfall
abundance or frequency. In southeast Australia mangroves are expanding as they migrate into
saltmarsh areas (Saintilan & Williams 1999, Harty & Cheng 2003, Harty 2004, Rogers et al. 2006).
At Botany Bay, New South Wales, mangrove area increased by 32.8% between 1956 and 1996
while saltmarsh coverage decreased by 78.7% (Evans & Williams 2001). Although no single factor
is responsible, it has been suggested that increased rainfall associated with climate change has
reduced salinity levels within salt marshes thereby allowing mangroves to migrate and outcompete
saltmarsh plants (Harty & Cheng 2003, Harty 2004, Rogers et al. 2006). However, hydrodynamic
modification related to urban and rural development is likely to be the overriding factor driving
mangrove expansion in the present climate.
Freshwater influx not only reduces the salinity of coastal waters but also enhances the stratifi-
cation of the water column thereby decreasing nutrient resupply from below. Flood events are
associated with an increase in productivity as nutrients are washed into the sea (McKinnon et al.
in press). While diatoms seem to be negatively affected by increases in river discharge, dinoflagel-
lates have been observed to profit from the increase in stratification and availability of humic
substances associated with riverine freshwater input (Carlsson et al. 1995, Goffart et al. 2002,
Edwards et al. 2006). Irrespective of the direction of change, modifications in rainwater runoff and
accompanying changes in salinity and resource supply should therefore affect the composition and,
potentially, the productivity of the phytoplankton community in coastal waters. River discharge is
also a primary shaper of soft-bottom coastal communities, particularly in tropical areas where
smaller watersheds produce more sediment (Rhoads et al. 1985, Milliman 1991, Alongi & Robert-
son 1995, Hall 2002).
Coral reefs are well known to be susceptible to fresh water as well as the effects of turbidity
and sedimentation that vary with coastal weather patterns. Numerous examples of coral commu-
nities being killed off or adversely affected purely as a result of extreme rainfall events have been
reported (Endean 1973, Rogers 1990, Alongi & Robertson 1995, Wilkinson 1999, Alongi &
McKinnon 2005, Fabricius 2005). Increases in rainfall, or extreme rainfall events, can increase
upland erosion and sediment transport considerably, thus severely impacting estuarine and near-
shore coastal ecosystems, especially along coastlines where development and other human land
uses have degraded the integrity of watersheds (Norkko et al. 2002, Thrush et al. 2003a,b, 2004,
444
2005, Lohrer et al. 2004). This is of particular concern for the Great Barrier Reef (Devlin & Brodie
2005).
Flood events are also associated with an increase in productivity as nutrients are washed into
the sea. These nutrients may also lead to undesirable effects, for example, producing ideal conditions
for the larvae of species such as the crown-of-thorns starfish. Recently long-term trends of increasing
nutrients and phytoplankton concentrations in the coastal waters of the Great Barrier Reef have
been shown, and these have been linked to recurring outbreaks of these starfish on the reef (Brodie
et al. 2005). These factors in combination appear to have resulted in large-scale trends in coral
diversity and abundance patterns on the Great Barrier Reef. Coastal areas in the so-called wet
tropics, characterised by higher rainfall, greater runoff, and the most intensive agriculture, have
generally lower coral diversity and lower coral cover than those adjacent to dryer coastal areas
(De Vantier et al. 2006).
Light penetration is an important factor limiting the distribution of marine macroalgae, both in
Australia (Kennelly 1989) and elsewhere (Reed & Foster 1984, Deysher & Dean 1986, Dayton
et al. 1999, Spalding et al. 2003). Increases in turbidity associated with greater rainfall, coastal
development or other human activities would thus be expected to degrade macroalgal communities
by generally decreasing the light penetration, so reducing depth ranges (Vadas & Steneck 1988;
also see review in Okey et al. 2004). Increases in turbidity can give competitive advantage to shade-
tolerant flora and non-photosynthetic organisms (Keough & Butler 1995). The ability of kelp to
compete with algal turfs may be reduced by coastal runoff. Turfs may benefit from the interaction
between sediment and nutrients (Gorgula & Connell 2004).
Cyclones and storms can be highly destructive by uprooting coastal plants, killing coastal
animals, and destabilising and eroding coastlines. These natural disturbance regimes may be
important in maintaining biodiversity in coastal ecosystems; however an increased frequency or
intensity of storms may reduce the resilience of coastal ecosystems (Dayton et al. 1992, Graham
1997, Carruthers et al. 2002, Fourqurean & Rutten 2004). Impacts on coral reefs can be severe;
the Category 4 Hurricane Andrew (Porter & Meier 1992) in Florida and other parts of the Caribbean
substantially damaged corals through intense wave impacts. Cyclones in Australia and elsewhere
have caused large-scale loss of algal cover and seagrass beds and devastation of mangroves and
coral reefs (Dayton et al. 1992, Preen et al. 1995, Rothlisberg et al. 1998, Gardner et al. 2005).
Recovery after these events can be relatively quick, but prolonged increases in the frequency or
intensity of storms and cyclones may increase the likelihood of severe perturbations and lead to
pronounced changes in biodiversity and community structure. Cyclones and storms also kill marine
animals such as turtles and seabirds and destroy breeding and feeding habitat (Limpus & Reed
1985). Destruction by cyclones is considered a major threat for breeding colonies of northern birds
such as the lesser noddy Anous tenuirostris melanops and the sooty tern Sterna fuscata (King et al.
1992, Garnett & Crowley 2000).
Storms also exert considerable damage in temperate ecosystems, for example through removal
of habitat-forming kelps and associated fauna, and influence community structure (Dayton & Tegner
1989). For example, the large fucoid alga (Carpophyllum flexuosum) is characteristic of calm
conditions and is now common in areas of northeastern New Zealand where it was once virtually
absent (Cole et al. 2001). This range expansion coincides with a significant decrease in storm
frequency and intensity in this part of New Zealand over the past 30 yr related to decadal-scale
climate variation (de Lange & Gibb 2000). Pinnipeds and seabirds nesting along Australia’s southern
shores are vulnerable to storm-induced mortality. Pup mortality in Australian fur seals is strongly
influenced by summer storms, particularly in low-lying colonies (Pemberton & Gales 2004).
Phenology The tropical wet season strongly influences life cycles of fauna and flora such as the
flowering and fruiting patterns of trees and shrubs (Friedel et al. 1993, Bach 2002, Keatley et al.
445
2002, Boulter et al. 2006) including mangroves (Ochieng & Erftemeijer 2002, Tyagi 2004). The
wet season also stimulates breeding in insects (Kemp 2001) and birds (Garnett & Crowley 2000,
Whitehead & Saalfeld 2000).
Tropical rainfall may also trigger behavioural changes in estuarine animals such as banana
prawns, Penaeus merguiensis. In common with many commercially important species of penaeid
prawns, these have a life cycle that involves migrations between nursery areas in mangrove-lined
creeks and estuaries and offshore coastal waters. Rainfall is highly correlated with offshore com-
mercial catches of banana prawns in southern areas of the Gulf of Carpentaria (Vance et al. 1985).
It is thought high rainfall leads to a decrease in salinity of estuarine waters, which triggers an
increased emigration of prawns (Staples 1980, Staples & Vance 1986, Vance et al. 1998). This
salinity trigger has been noted in other parts of the world (see Zein-Eldin & Renaud 1986).
Pelagic systems
Physiology Because coastal regions may receive considerable freshwater input, coastal phy-
toplankton is subject to more variation in salinity than oceanic phytoplankton. In general phyto-
plankton species are adapted for ambient salinity so coastal species such as Skeletonema costatum
thrive over a wide range of salinities while offshore species grow well only within narrow salinity
ranges. In addition, estuarine and coastal species exhibit optimal growth at low and intermediate
salinities while offshore species thrive at high salinities (Brand 1984, McQuoid 2005, Thessen et al.
2005).
Sea level
Coastal systems
Physiology Different mangrove species are adapted for different tidal inundation regimes as
apparent in the zonation patterns of coastal mangroves. Rising sea level will alter the tidal inundation
regime experienced by mangroves and presumably increase environmental stress on individual
plants. For example, laboratory experiments have shown increased tidal inundation reduced growth
and photosynthetic rates in Rhizophora mangle seedlings (Ellison & Farnsworth 1997). At any
particular site, the mangrove community is highly specialised for local environmental conditions
so minor variations in hydrological or tidal regimes can result in high mortality (Blasco et al. 1996).
Distribution and abundance Coastal marine habitats will be vulnerable to changes in sea level
(Short & Neckles 1999). The increase in water depth and consequent reduction in light availability
to the sea bed will impact subtidal marine plants and tropical corals. At any given location, the
location of maximum depth limits will shift, depending on topography, directly affecting distribu-
tions and abundance. For example, it is estimated that a 50-cm increase in sea level could result
in a 30–40% reduction in growth of Zostera marina, a widespread Northern Hemisphere seagrass
(Short & Neckles 1999). In many places, the shoreward shift of plants and animals will be impeded
by coastal development.
Sea-level rise will also alter the magnitude of local tidal ranges, depending on interactions with
coastal topography. An increase or decrease in tidal range will directly impact the ‘zonation’ of
macroalgae and fauna in the intertidal and subtidal. An increase in tidal range will exacerbate
effects of changing water depth on subtidal plant communities, resulting in a loss of biomass in
deeper waters whereas a decrease in tidal range will reduce exposure stress at shallower depths
(Short & Neckles 1999).
Mangroves typically occur on low-profile, low-energy coastlines and are ecologically restricted
to saline intertidal environments so are considered particularly susceptible to rapid changes in sea
446
level (Woodroffe 1992, Ellison 1993, Parkinson et al. 1994, Field 1995). Mangrove areas around
Australia with small tidal regimes such as Shark Bay and the Exmouth Gulf are likely to be
inundated by the projected rise in sea level (see Hughes 2003). When sea levels rise, as projected
over the next century, shorelines will move landward but if sedimentation is more rapid then sea-
level rise, shorelines may actually move seaward. Mangroves trap suspended sediments, for example
a field study in a mangrove swamp in Cairns found 80% of suspended sediment brought in from
coastal waters at spring flood tide was trapped in the mangroves, resulting in a rise of the substratum
of 1 mm per year and presumably reducing turbidity in coastal waters (Furukawa et al. 1997).
If sediment accretion rates in mangroves are equal to or exceed sea-level rise then mangroves
will persist. The ability to accrete sediments will depend on the availability of suspended sediments
in coastal waters so in areas where suspended sediment load is low, mangroves may not be able
to track rising sea levels (Ellison & Stoddart 1991, Parkinson et al. 1994). Modelling studies of
the response of tropical Australian estuaries to sea-level rise reveal differing responses depending
on hydrodynamics and channel morphology; in some estuaries mangroves will expand while in
others mangroves will retreat (Wolanski & Chappell 1996). Mangroves growing on carbonate
settings or low islands may be strongly threatened by sea-level rise over the next century (Ellison &
Stoddart 1991, Ellison 1993).
Rising sea level is also a threat to bird species that nest on low-lying coastal areas as valuable
breeding sites are flooded or eroded (Galbraith et al. 2002). Examples are the little kingfisher Alcedo
pusilla pusilla and spangled drongo Dicrurus bracteatus carbonarius nesting on low-lying islands
in the Torres Strait and the lesser noddy Anous tenuirostris melanops that nests in mangroves
(Garnett & Crowley 2000). Marine turtle breeding beaches will be impacted by sea-level rise. For
example, 32% of current beach area on the island of Bonaire in the Caribbean could be lost if the
sea level rises by 50 cm and the loss of potential turtle nesting habitat may be even higher particularly
where land directly behind the beach system is developed (Fish et al. 2005). Pinniped haul-out sites
for breeding and nurseries may also be reduced or eliminated by sea-level rise (Learmonth et al.
2006).
Community impacts
Climate impacts on particular species or groups do not occur in isolation and can result in extensive
cascading effects and complex interactions (Figure 8). Climatic impacts on a few leverage species,
such as species considered foundation species or ecosystem engineers, may result in sweeping
community-level changes (Coleman & Williams 2002). Foundation species such as corals support
a diverse range of fauna and flora by providing complex architectures of living habitat while
ecosystem engineers increase habitat complexity either morphologically or behaviourally. Species
that are functionally unique play a distinct role, so loss of these species tends to result in severe
impacts on the ecosystem (Fonseca & Ganade 2001). Others such as phytoplankton and zooplankton
are found in great numbers and are the base of trophic webs. Most of these groups are primary or
secondary producers and therefore support higher trophic levels such as pelagic and demersal fishes,
seabirds, turtles and marine mammals. Keystone species have a disproportionate structuring effect
on biological communities (large interaction strength relative to their own abundance or biomass),
and they are often vulnerable to local extinction due to their small numbers or biomass.
Models and analytical tools provide the capability to estimate climate change impacts in terms
of diversity, community composition and species interactions in the context of both direct and
indirect effects. Relatively little modelling work has been done on Australian marine species and
communities, and there has been no large-scale investigation on the potential impacts of climate
change on the diverse and unique fauna of the region. Corals of the Great Barrier Reef are the only
group that has been investigated extensively in terms of potential impacts of climate change.
447
Stressors
Climate change
(e.g. increased temperatures, increased storms,
Low currents, acidification) High
Non-climate change
(e.g. fisheries, pollution, habitat degradation)
Attributes of species that arrive or leave
Net interaction strength

Engineering/architecture-forming (foundational)
Low High
Keystoneness
Functional uniqueness
Small change Community structure Large change
Management strategies
Mitigation by global emissions reductions
Major Adaptation by management of Minor
non-climate stressors
Figure 8 Species attributes, types of stressors and management strategies that influence the magnitude of
climate change impacts on the structure and function of biological communities. Low levels of particular
attributes of the species that invade an area or become locally extinct lead to minimal changes, as do low
levels of climate and non-climate stressors that a community is exposed to. High levels of those attributes and
stressors lead to large community changes. Major implementation of management strategies can reduce
community impacts, whereas minor implementation in the context of major stressors can lead to large
community changes.
Non-climate stressors
Climate change is not the only stressor to impact ecosystems, either at present or into the future.
Anthropogenic stressors, such as fishing, pollution, coastal development and exotic pests, will all
decrease the resilience of marine life and ecosystems. Systems that are already highly stressed may
be particularly vulnerable to further perturbations such as those induced by climate change (Hughes
& Connell 1999, Steneck et al. 2002, Hughes et al. 2003). Most non-climate stressors can be
managed faster than climate change by altering policy and management practices on national and
regional scales.
Although Australian fisheries are relatively small by international standards due to the generally
oligotrophic waters, considerable tonnage is still extracted and many species and groups are
overexploited or at high risk. Fisheries can have major impacts on marine systems through removal
of large predators, substantial by-catch of non-target species and habitat destruction by dredges
and trawls. Australian fisheries include various commercial (Caton & McLaughlin 2004), recre-
ational and indigenous fisheries (Henry & Lyle 2003). Commercial fisheries harvest more than
130,000 tonnes of fish, squid and crustaceans annually, mostly from coastal, continental shelf and
upper continental slope waters. Of 74 Australian stocks considered in 2004, 17 were overfished,
17 were not overfished and 40 were of uncertain status (Caton & McLaughlin 2004), but such
448
statistics can underestimate the damage to non-target biota and habitat structure. Recreational fishers
also remove a sizeable biomass of fish and crustaceans (>30,000 tonnes annually) from coastal and
estuarine waters (Lyle et al. 2003).
Virtually all of Australia’s population and industries are situated along the coastal fringe or
rivers that drain into the sea, and so the effects of pollution and coastal development on marine
species or ecosystems can be severe (Kirkman 1997, Duke et al. 2005, Votier et al. 2005). Modi-
fication of structure and function of coastal watersheds by agriculture, urban development, and
deforestation can lead to considerable increases in erosion and nutrient runoff. Habitat modification
and destruction through coastal development or activities such as dredging will all impact marine
habitats such as estuaries, mangroves, seagrass beds and kelp forests. These habitats are integral
features of marine ecosystems that provide a variety of critical ecosystem services such as nursery
grounds, primary production and adult habitats for whole suites of marine organisms.
Introduced species can also have severe consequences for marine ecosystems. For example in
1995 and 1998/1999, mass mortalities linked to exotic pathogens probably introduced from aqua-
culture feed spread rapidly throughout the Australian population of the sardine Sardinops sagax
(Gaughan 2001, Ward et al. 2001). These mortality events represent two of the most extensive mass
mortality events recorded for marine organisms (Gaughan 2001).
Biological communities have adapted to various levels of natural disturbance and variability
over evolutionary timescales. Shifts in these disturbance regimes increase stress to these systems
and decrease the overall resilience of the system to other disturbances. However, the resilience
principle also implies that reductions of the stressors that humans can control may partially
ameliorate increasing climate change impacts (Figure 8). Thus easing the impacts of fisheries,
pollution, habitat destruction and other non-climate anthropogenic-induced stressors on marine
ecosystems may partly mitigate climate change impacts. Although immediate and conscientious
international diplomacy to reduce greenhouse gas emissions is a critical mitigation strategy for
addressing the long-term impacts of climate changes, adaptive and integrated management systems
that focus on fisheries and pollution on regional levels are just as critical because these can address
the near-term inevitable changes that will act synergistically with climate change to threaten
Australia’s marine life.
Summary
Rising temperatures will have a major influence on species distributions, although population
responses will be modified by climate-induced changes in competitive ability, dispersive capacity
and behaviour of organisms. However, a general shift in species distributions toward higher latitudes
is expected and is already occurring in many parts of the world (Parmesan et al. 1999, Thomas &
Lennon 1999, Beaugrand et al. 2002, Parmesan & Yohe 2003, Hickling et al. 2006). In Australian
coastal waters, this shift may be facilitated by the major southward-flowing surface currents,
particularly given the projected enhancement of the EAC. A concurrent alteration in phenology is
expected, with longer growing seasons for marine plants (e.g., seagrasses) and earlier breeding
seasons of marine animals. Higher sea level will alter coastline and island hydrography and
topography, with potential loss of nesting or breeding areas for seabirds, turtles and seals. Acidi-
fication may become a major threat to tropical coral reefs and the cold-water corals found on the
edge of the continental slope and on seamounts and to some plankton that are important for
ecosystem functioning (Orr et al. 2005, Guinotte et al. 2006).
A schematic of many of the expected impacts of future climate change on Australian marine
systems is shown in Figure 9. In coastal waters, tropical species of seagrasses, mangroves and fish
have shifted further south. Dugongs have also moved further south following the expansion of
tropical seagrasses. However, cold-water kelp species have disappeared from higher latitudes with
449
Figure 9 (See also Colour Figure 9 in the insert.) Hypothetical Southern Hemisphere marine coastline and
coastal waters ranging from low latitudes in the north to high latitudes in the south under present climate
(top) and in the future under global warming scenario (bottom). As temperatures rise, species’ distributions
shift further south. The range of tropical and subtropical species extends to temperate latitudes while temperate
species in the south decline. Rising temperatures and ocean acidification stress coral reefs leading to frequent
coral bleaching and an increase in mortality while rapid sea level rise inundates the coral reefs. Ocean
acidification also leads to the decline of calcifying plankton such as pteropods and coccolithophores. Rising
sea-level encroaches on the mainland and on offshore islands. The sex ratio of marine turtle hatchlings, which
is determined by ambient nest temperatures, is skewed in the future as warming produces more females.
450
a warming climate and the occurrence and distribution of venomous jellyfish has increased. Further
offshore corals have bleached and declined in response to warmer temperatures and ocean acidifi-
cation. Tropical pteropods and coccolithophores have also declined as the oceans acidify. The sex
ratio of turtle hatchlings is heavily skewed toward females as nesting beaches heat up. Finally,
rapid sea-level rise has drowned the coastline, islands and barrier reefs. Monitoring and strategic
planning is sorely needed for Australia because numerous other climate change effects might be
already occurring in Australia’s marine realm.
Tropical and subtropical Australia

Rising temperatures and ocean acidification are considered the major climate change threats to
tropical coral reefs. Bleaching of coral reefs has occurred regularly over the last couple of decades
and is projected to occur with increasing frequency over this century. Australian coral reefs are
relatively healthy compared to many elsewhere in the world (Pandolfi et al. 2005). However, they
are facing considerable pressures from changes to coastal water quality and overexploitation of key
ecological functional groups, particularly in areas close to urban and agricultural developments.
The role of coral reefs in underpinning coastal economies in Australia is becoming increasingly
recognised. The pristine nature of Australian coral reefs attracts large numbers of tourists; the reef-
associated tourist economy currently exceeds $A4 billion per annum. Hoegh-Guldberg & Hoegh-
Guldberg (2004) analysed the potential effect of losing the competitive edge for tourism if the coral
reefs on the Great Barrier Reef deteriorated as a result of climate change. The effects vary, depending
on domestic and international trends in aspects such as markets and politics. However, if one uses
the reef-associated component of these economies as a guide to how things might change if reefs
continue to degrade, degradation would reduce international tourist income by as much as $A8
billion over 19 yr (Hoegh-Guldberg & Hoegh-Guldberg 2004).
Seagrass beds and mangroves commonly co-occur with tropical coral reefs and there are strong
interactions between them (Harborne et al. 2006). Coral reef crests dissipate wave energy thus
ensuring the calm conditions required by seagrass beds and mangroves further inshore, while
mangrove and seagrasses filter riverine sediments and nutrients that would otherwise impact coral
reefs. Many fish are found in mangroves and seagrasses as juveniles before undertaking ontogenetic
shifts in habitat use onto coral reefs (Mumby et al. 2004, Dorenbosch et al. 2005, Mumby 2006).
Deterioration in one or more of these systems due to climate change or other impacts may have
deleterious consequences for the entire coastal ecosystem with associated economic losses. A
reduction in seagrass beds and mangroves will have an immediate impact on the economic value
of associated fisheries (Loneragan et al. 2005, Manson et al. 2005, McArthur & Boland 2006).
Mangroves and seagrasses provide a range of ecosystem services such as recycling of carbon and
nutrients, shoreline protection and enriched coastal productivity (Costanza et al. 1997, Ewel et al.
1998, Kathiresan & Bingham 2001, Duarte et al. 2005, Bloomfield & Gillanders 2005). Coastal
fringing mangroves are important for shoreline protection from storms and erosion (Ewel et al.
1998, Badola & Hussain 2005, Kathiresan & Rajendran 2005).
Temperate Australia
Over 46% of Australian’s population live in the southeast (Zann 2000). Around Sydney, central
New South Wales, the coastline is largely metropolitan with extensive industrial development (Zann
2000). The southeast is considered the most stressed from anthropogenic pressures, other than
climate change, such as metal and sewage pollution (Hobday et al. 2006). Large demersal trawl
fisheries operate in southeastern Australian waters. This region is also considered the most stressed
451
by fishing pressure, with a highest proportion of overexploited stocks (Caton & McLoughlin 2004,
Hobday et al. 2006).
Marine ecosystems of southern Australia are strongly influenced by the Leeuwin Current and
EAC (Maxwell & Cresswell 1981, Phillips 2001). The temperature difference between the EAC
and surrounding waters can be more than 5°C (Zann 2000). The projected strengthening of the
EAC and warming of the Tasman Sea as global climate warms will have detrimental effects for
cold-temperate species in the southeast, particularly in Tasmanian waters. The shelf does not extend
far south of Australia and the lack of alternative land mass until Antarctica means these species
have no suitable habitat to occupy as global climate warms. The cold-water giant kelp is already
in decline in this region and presumably other marine organisms in this region are also at high risk
from climate change.
All marine groups investigated are expected to show some southward movement of members.
The coccolithophore Gephyrocapsa oceanica is a good example; a tropical strain of it has already
expanded into temperate waters in eastern Australia. The relatively productive temperate pelagic
zone may shrink considerably in area and potentially become restricted to west of Tasmania by the
2070s, while the food web in formerly productive regions may shift toward a much less productive
subtropical regime. The recent expansion of G. oceanica and decreasing stock sizes of jack mackerel
in southeastern shelf waters and southern bluefin tuna returning to the east coast in winter indicate
that the pelagic ecosystem may have started to change (Welsford & Lyle 2003, Blackburn 2005,
Polacheck et al. 2006).
Venomous jellyfish and harmful algal blooms are a major threat to human health, but are largely
phenomena associated with tropical waters or relatively warm and stratified waters, respectively.
As a consequence of enhanced southward flow of warm currents such as the EAC and ocean
warming these phenomena will likely extend into more southerly regions currently unaffected
(jellyfish) or occur more frequently (in case of harmful algal blooms).
Critical knowledge gaps and a way forward

In this review we have detailed the expected considerable and observed consequences of climate
change for marine groups, some of which, such as bleaching of tropical corals, are already being
observed in Australian waters. At present, it is impossible to determine if climate change is
impacting many less-charismatic marine groups and habitats in Australia, despite compelling
evidence from elsewhere in the world.
Long-term datasets are key to documenting and understanding the response of species to climate
change. Australian marine scientists have long claimed that the lack of observable climate signals
is a consequence of the paucity of ecological time series in the region. This claim is not unique to
Australia. Despite an exponential increase in the initiation of long-term monitoring programmes
in the world’s oceans since World War II, 40% of these time series were discontinued during the
1980s because monitoring was viewed as poor science by administrators (Duarte et al. 1992). This
negative perception began shifting during the late 1990s when the knowledge of consequences of
climate change began emerging in scientific and political realms, and this has markedly improved
the support for many monitoring programmes (Hays et al. 2005).
The case of zooplankton sampling in Australia highlights gaps in the present monitoring system.
Zooplankton may be the most abundant multicellular organisms on the planet, are the major source
of food of many marine organisms, and are considered sentinels of climate change (Hays et al.
2005). Globally there are zooplankton time series spanning more than 15 yr in no fewer than 30
countries, including relatively small or developing nations such as Bulgaria, Chile, Estonia, Greece,
Kazakhstan, Latvia, Faroe Islands, Namibia, Peru, Turkey and the Ukraine. However, the longest
452
ongoing zooplankton time series in Australia is 2 yr and consists of a single cross-shelf transect
off Perth. Given the diversity of marine habitats in Australia and the economic and social importance
of fishing, Australia is clearly impoverished in long-term zooplankton and other datasets urgently
required to assess climate change impacts (see Hobday et al. 2006). Without such datasets, Australia
will be unaware of how its marine systems are altered by future climate change, continuing to rely
on information gathered from systems elsewhere. This will make adaptation and mitigation strat-
egies uncertain.
This review indicates that we have a general understanding of some of the likely mechanisms
of climate effects on a few particular species, but we have limited knowledge about how Australian
marine ecosystems will respond to climate change. It is only when Australia focuses on the entirety
of its marine resources will we be able to tackle rigorously the impacts of climate change. There
are a number of critical questions that need to be addressed to allow managers tasked with
conserving biodiversity, locating marine protected areas, managing eco-tourism associated with
cetaceans and turtles, and implementing management plans for the sustainable use of marine
resources and indigenous harvesting:
• How will the distribution, abundance and phenology of marine species alter with climate
change and how will these impact communities?
• Which species are candidate indicators to monitor climate change in Australian waters?
• Which areas are particularly sensitive to changing climate or are ‘hot spots’ of change?
• How will regional ocean productivity alter with climate change?
• How can ecosystem resilience to climate change be increased?
• How will climate change affect the socioeconomic productivity of marine ecosystems?
Acknowledgements
This contribution was supported by the CSIRO Wealth from Oceans National Research Flagship
and the Australian Greenhouse Office.
References
Abal, E.G., Loneragan, N., Bowen, P., Perry, C.J., Udy, J.W. & Dennison, W.C. 1994. Physiological and
morphological responses of the seagrass Zostera capricorni Aschers to light intensity. Journal of
Adey, W.H. & Steneck, R.S. 2001. Thermography over time creates biogeographic regions: a tempera-
ture/space/time-integrated model and an abundance-weighted test for benthic marine algae. Journal
of Phycology 37, 677–698.
Adnyana, W., Ladds, P.W. & Blair, D. 1997. Observations of fibropapillomatosis in green turtles (Chelonia
mydas) in Indonesia. Australian Veterinary Journal 75, 737–742.
Ainsworth, E.A. & Long, S.P. 2005. What have we learned from 15 years of Free Air Carbon Dioxide
Enrichment (FACE)? A meta-analytic review of the responses of photosynthesis, canopy properties
and plant production to rising CO2. New Phytologist 165, 351–371.
Alongi, D.M. & McKinnon, A.D. 2005. The cycling and fate of terrestrially-derived sediments and nutrients
in the coastal zone of the Great Barrier Reef shelf. Marine Pollution Bulletin 51, 239–252.
Alongi, D.M. & Robertson, A.I. 1995. Factors regulating benthic food chains in tropical river deltas and
adjacent shelf areas. Geo-Marine Letters 15, 145–152.
Andrew, N.L. 1991. Changes in subtidal habitat following mass mortality of sea urchins in Botany Bay, New
South Wales. Australian Journal of Ecology 16, 353–362.
453
Antia, A.N., Koeve, W., Fischer, G., Blanz, T., Schul-Bull, D., Scholten, J., Neuer, S., Kremling, K., Kuss, J.,
Peinert, R., Hebbeln, D., Bathmann, U., Conte, M. & Fehner, U. 2001. Basin-wide particulate carbon
flux in the Atlantic Ocean: regional export patterns and potential for atmospheric CO2 sequestration.
Global Biogeochemical Cycles 15, 845–862.
Archaux, F. 2003. Birds and climate change. Vie et Milieu 53, 33–41.
Arsenault, D.J., Marchinko, K.B. & Palmer, A.R. 2001 Precise tuning of barnacle leg length to coastal wave
action. Proceedings of the Royal Society of London B 268, 2149–2154.
Australia Bureau of Statistics. 2006. 2001 Census Data. Available HTTP: http://www.abs.gov.au (accessed
10 July 2006).
Australian Greenhouse Office. 2003. Climate Change: An Australian Guide to the Science and Potential
Impacts, B. Pittock (ed.). Canberra: Australian Greenhouse Office.
Babcock, R. & Davies, P. 1991. Effects of sedimentation on settlement of Acropora millepora. Coral Reefs
9, 205–208.
Babcock, R.C., Bull, G.D., Harrison, P.L., Heyward, A.J., Oliver, J.K., Wallace, C.C. & Willis, B.L. 1986.
Synchronous spawning of 105 scleractinian coral species on the Great Barrier Reef. Marine Biology
90, 379–394.
Babcock, R.C. & Smith, L. 2002. Effects of sedimentation on coral settlement and survivorship. In Proceedings
of the Ninth International Coral Reef Symposium, Bali, Indonesia, October 23–27, 2000, M.K. Kasim
Moosa et al. (eds). Jakarta, Indonesia: Ministry of Environment, the Indonesian Institute of Sciences
and the International Society for Reef Studies, 245–248.
Babcock, R.C., Willis, B.L. & Simpson, C.J. 1994. Mass spawning of corals on a high latitude coral reef.
Coral Reefs 13, 161–169.
Bach, C.S. 2002. Phenological patterns in monsoon rainforests in the Northern Territory, Australia. Austral
Ecology 27, 477–489.
Badola, R. & Hussain, S.A. 2005. Valuing ecosystem functions: an empirical study on the storm protection
function of Bhitarkanika mangrove ecosystem, India. Environmental Conservation 32, 85–92.
Bancroft, W.J., Garkaklis, M.J. & Roberts, J.D. 2004. Continued expansion of the wedge-tailed shearwater,
Puffinus pacificus, nesting colonies on Rottnest Island, Western Australia. Emu 104, 79–82.
Barbieri, E.S., Villafane, V.E. & Helbling, W. 2002. Experimental assessment of UV effects on temperate
marine phytoplankton when exposed to variable radiation regimes. Limnology and Oceanography 47,
1648–1655.
Barnes, J.H. 1964. Cause and effect in Irukandji stingings. Medical Journal of Australia 14, 897–904.
Barry, J.P., Baxter, C.H., Sagarin, R.D. & Gilman, S.E. 1995. Climate-related, long-term faunal changes in a
California rocky intertidal community. Science 267, 672–675.
Baruch, R., Avishai, N. & Rabinowitz, C. 2005. UV incites diverse levels of DNA breaks in different cellular
compartments of a branching coral species. Journal of Experimental Biology 208, 843–848.
Beamish, R.J., Mahnken, C. & Neville, C.M. 1997. Hatchery and wild production of Pacific salmon in relation
to large-scale, natural shifts in the productivity of the marine environment. ICES Journal of Marine
Science 54, 1200–1215.
Beamish, R.J., Noakes, D.J., McFarlane, G.A., Klyashtorin, L., Ivanov, V.V. & Kurashov, V. 1999. The regime
concept and natural trends in the production of Pacific salmon. Canadian Journal of Fisheries and
Aquatic Sciences, 56, 516–526.
Beardall, J., Beer, S. & Raven, J.A. 1998. Biodiversity of marine plants in an era of climate change: some
predictions based on physiological performance. Botanica Marina 41, 113–123.
Beardall, J. & Raven, J.A. 2004. The potential effects of global climate change on microalgal photosynthesis,
growth and ecology. Phycologia 43, 26–40.
Beare, D.J., Burns, F., Greig, A., Jones, E.G., Peach, K., Kienzle, M., McKenzie, E. & Reid, D.G. 2004. Long-
term increases in prevalence of North Sea fishes having southern biogeographic affinities. Marine
Beaugrand, G., Reid, P.C., Ibanez, F., Lindley, J.A. & Edwards, M. 2002. Reorganisation of North Atlantic
marine copepod biodiversity and climate. Science 296, 1692–1694.
Beaumont, L.J., McAllan, I.A. & Hughes, L. 2006. A matter of timing: changes in the first date of arrival and
last date of departure of Australian migratory birds. Global Change Biology 12, 1339–1354.
454
Behrenfeld, M., Hardy, J., Gucinski, H., Hannemann, A., Lee, H., II & Wones, A. 1993. Effects of ultraviolet-
B radiation on primary production in the South Pacific Ocean. Marine Environmental Research 35,
349–363.
Berge, J.A., Bjerkeng, B., Pettersen, O., Schaanning, M.T. & Oxnevad, S. 2006. Effects of increased sea water
concentrations of CO2 on growth of the bivalve Mytilus edulis L. Chemosphere 62, 681–687.
Binckley, C.A., Spotila, J.R., Wilson, K.S. & Paladino, F.V. 1998. Sex determination and sex ratios of Pacific
leatherback turtles, Dermochelys coriacea. Copeia 2, 291–300.
Bischof, K., Hanelt, D. & Wiencke, C. 1998. UV-radiation can affect depth-zonation of Antarctic macroalgae.
Blackburn, S. 2005. Coccolithophorid species and morphotypes — indicators of climate change. In Regional
Impacts of Climate Change and Variability in South-East Australia, V. Lyne et al. (workshop conve-
nors). Hobart, Australia: CSIRO Marine Research and CSIRO Atmospheric Research, 14–15.
Blackburn, S.I. & Cresswell, G. 1993. A coccolithophorid bloom in Jervis Bay, Australia. Australian Journal
of Marine and Freshwater Research 44, 785–786.
Blackwell, B. 2005. The economic value of Australia’s natural coastal assets: some preliminary findings.
Ecological Economics in Action Conference, December 11–13 2005, Massey University, Palmerston
North, New Zealand, 143–151. Available HTTP: http://www.anzsee.org/decconferindex.asp (accessed
28 September 2006).
Blasco, F., Saenger, P. & Janodet, E. 1996. Mangroves as indicators of coastal change. Catena 27, 167–178.
Bloomfield, A.L. & Gillanders, B.M. 2005. Fish and invertebrate assemblages in seagrass, mangrove, salt-
marsh, and nonvegetated habitats. Estuaries 28, 63–77.
Bollens, S.M. & Frost, B.W. 1990. UV light and vertical distribution of the marine planktonic copepod Acartia
hudsonica Pinhey. Journal of Experimental Marine Biology and Ecology 137, 89–93.
Bolton, J.J. 1996. Patterns of species diversity and endemism in comparable temperate brown algal floras.
Hydrobiologia 327, 173–178.
Bonaventura, R., Poma, V., Russo, R., Zito, F. & Matranga, V. 2006. Effects of UV-B radiation on development
and hsp70 expression in sea urchin cleavage embryos. Marine Biology 149, 79–86.
Bonnet, D., Richardson, A.J., Harris, R., Hirst, A., Beaugrand, G., Edwards, M., Ceballos, S., Diekman, R.,
López-Urrutia, A., Valdes, L., Carlotti, F., Molinero, J.C., Weikert, H., Greve, W., Lucic, D., Albaina,
A., Yahia, N.D., Umani, S.F., Miranda, A., dos Santos, A., Cook, K., Robinson, S. & Fernandez de
Puelles, M.L. 2005. An overview of Calanus helgolandicus ecology in European waters. Progress in
Booth, D.T. & Astill, K. 2001. Temperature variation within and between nests of the green sea turtle, Chelonia
mydas (Chelonia: Cheloniidae) on Heron Island, Great Barrier Reef. Australian Journal of Zoology
49, 71–84.
Bopp, L., Monfray, P., Aumont, O., Dufresne, J.L., Le Treut, H., Madec, G., Terreay, L. & Orr, J.C. 2001.
Potential impact of climate change on marine export production. Global Biogeochemical Cycles 15,
81–99.
Both, C., Artemyev, A.V., Blaauw, B., Cowie, R.J., Dekhuijzen, A.J., Eeva, T., Enemar, A., Gustafsson, L.,
Ivankina, E.V., Jarvinen, A., Metcalfe, N.B., Nyholm, N.E.I., Potti, J., Ravussin, P.A., Sanz, J.J.,
Silverin, B., Slater, F.M., Sokolov, L.V., Torok, J., Winkel, W., Wright, J., Zang, H. & Visser, M.E.
2004. Large-scale geographical variation confirms that climate change causes birds to lay earlier.
Proceedings of the Royal Society of London Series B 271, 1657–1662.
Both, C., Piersma, T. & Roodbergen, S.P. 2005. Climatic change explains much of the twentieth century
advance in laying date of Northern Lapwing Vanellus vanellus in the Netherlands. Ardea 93, 79–88.
Boulding, E.G., Holst, M. & Pilon, V. 1999. Changes in selection on gastropod shell size and thickness with
wave-exposure on northeastern pacific shores. Journal of Experimental Marine Biology and Ecology
232, 217–239.
Boulter, S.L., Kitching, R.L. & Howlett, B.G. 2006. Family, visitors and the weather: patterns of flowering
in tropical rain forests of northern Australia. Journal of Ecology 94, 369–382.
Bowen, B.W., Abreu-Grobois, F.A., Balazs, G.H., Kamezaki, N., Limpus, C.J. & Ferl, R.J. 1995. Trans-Pacific
migrations of the loggerhead turtle (Caretta caretta) demonstrated with mitochondrial DNA markers.
Proceedings of the National Academy of Sciences of the United States of America 92, 3731–3734.
455
Bowman, D.M.J.S., Walsh, A. & Milne, D.J. 2001. Forest expansion and grassland contraction with a Euca-
lyptus savanna matrix between 1941 and 1994 at Litchfield National Park in the Australian monsoon
tropics. Global Ecology and Biogeography 10, 535–548.
Boyd, P.W. & Doney, S.C. 2002. Modelling regional responses by marine pelagic ecosystems to global climate
change. Geophysical Research Letters 29, 1806.
Boyd, P.W., Watson, A.J., Law, C.S., Abraham, E.R., Trull, T., Murdoch, R., Bakker, D.C.E., Bowie, A.R.,
Buesseler, K.O., Chang, H., Charette, M., Croot, P., Downing, K., Frew, R., Gall, M., Hadfield, M.,
Hall, J., Harvey, M., Jameson, G., LaRoche, J., Liddicoat, M., Ling, R., Maldonado, M.T., McKay,
R.M., Nodder, S., Pickmere, S., Pridmore, R., Rintoul, S., Safi, K., Sutton, P., Strzepek, R., Tan-
neberger, K., Turner, S., Waite, A. & Zeldis, J. 2000. A mesoscale phytoplankton bloom in the polar
Southern Ocean stimulated by iron fertilization. Nature 407, 695–702.
Brand, L.E. 1984. The salinity tolerance of 46 marine phytoplankton isolates. Estuarine, Coastal and Shelf
Science 18, 543–556.
Brawley, S.H. 1992. Fertilization in natural populations of the dioecious brown alga Fucus ceranoides and
the importance of polyspermy blocks. Marine Biology 113, 145–157.
Brawley, S.H., Johnson, L.E., Pearson, G.A., Speransky, V., Li, R. & Serrão, E. 1999. Gamete release at low
tide in fucoid algae: maladaptive or advantageous? American Zoologist 39, 218–229.
Broderick, A.C., Godley, B.J. & Hays, G.C. 2001. Trophic status drives interannual variability in nesting
numbers of marine turtles. Proceedings of the Royal Society of London Series B 268, 1481–1487.
Brodie, J., Fabricius, K., De’ath, G. & Okaji, K. 2005. Are increased nutrient inputs responsible for more
outbreaks of crown-of-thorns starfish? An appraisal of the evidence. Marine Pollution Bulletin 51,
266–278.
Bruhn, J. & Gerard, V.A. 1996. Photoinhibition and recovery of the kelp Laminaria saccharina at optimal
and superoptimal temperatures. Marine Biology 125, 639–648.
Bunce, A., Norman, F.I., Brothers, N. & Gales, R. 2002. Long-term trends in the Australasian gannet (Morus
serrator) population in Australia: the effect of climate change and commercial fisheries. Marine
Biology 141, 263–269.
Burkett, V., Codignotto, J.O., Forbes, D. L., Mimura, N., Beamish, R.J. & Ittekkot, V. 2001. Marine Fish. In
Climate 2001: Coastal Zones and Marine Ecosystems. Contribution of Working Group II to the Third
Assessment Report of the Intergovernmental Panel on Climate Change. J.J. McCarthy et al. (eds)
Cambridge, UK & New York, USA: Cambridge University Press, 350-353
Byrkjedal, I., Godo, O.R. & Heino, M. 2004. Northward range extensions of some mesopelagic fishes in the
Northeastern Atlantic. Sarsia 89, 484–489
Cai, W. 2006. Antarctic ozone depletion causes an intensification of the Southern Ocean super-gyre circulation.
Geophysical Research Letters 33, L03712.
Cai, W., Shi, G., Cowan, T., Bi, D. & Ribbe, J. 2005. The response of the Southern Annular Mode, the East
Australian Current, and the southern mid-latitude ocean circulation to global warming. Geophysical
Research Letters 32, L23706.
Caldeira, K. & Wickett, M.E. 2003. Anthropogenic carbon and ocean pH. Nature 425, 365–365.
Caldeira, K. & Wickett, M.E. 2005. Ocean model predictions of chemistry changes from carbon dioxide
emissions to the atmosphere and ocean. Journal of Geophysical Research — Oceans 110, C09S04.
Cambridge, M.L. & Hocking, P.J. 1997. Annual primary production and nutrient dynamics of the seagrasses
Posidonia sinuosa and Posidonia australis in south-western Australia. Aquatic Botany 59, 277–295.
Campbell, R.A. & Hobday, A. 2003. Swordfish–environment–seamount–fishery interactions off eastern Aus-
tralia. Report to the Australian Fisheries Management Authority, Canberra, Australia.
Campbell, S.J., McKenzie, L.J. & Kerville, S.P. 2006. Photosynthetic responses of seven tropical seagrasses
to elevated seawater temperature. Journal of Experimental Marine Biology and Ecology 330, 455–468.
Caputi, N., Chubb, C. & Pearce, A. 2001. Environmental effects on recruitment of the western rock lobster,
Panulirus cygnus. Marine and Freshwater Research 52, 1167–1174.
Carlsson, P., Graneli, E., Tester, P. & Boni, L. 1995. Influences of riverine humic substances on bacteria,
protozoa, phytoplankton, and copepods in a coastal plankton community. Marine Ecology Progress
Series 127, 213–221.
456
Carr, A. 1987. New perspectives on the pelagic stage of sea turtle development. Conservation Biology 1,
103–121.
Carruthers, T.J.B., Dennison, W.C., Longstaff, B.J., Waycott, M., Abal, E.G., McKenzie, L.J. & Lee Long,
W.J. 2002. Seagrass habitats of northeast Australia, models of key processes and controls. Bulletin of
Marine Science 71, 1153–1170.
Caton, A. & McLoughlin, K. (eds) 2004. Fishery Status Reports 2004: Status of Fish Stocks Managed by the
Australian Government. Canberra, Australia: Bureau of Rural Sciences.
Chaloupka, M. 2001. Historical trends, seasonality and spatial synchrony in green sea turtle egg production.
Biological Conservation 101, 263–279.
Chambers, L.E. 2004. The Impact of Climate on Little Penguin Breeding Success. BMRC Research Report
100. Melbourne, Australia: Bureau of Meteorology.
Chambers, L.E. 2006. Associations between climate change and natural systems in Australia. Bulletin of the
American Meteorological Society 87, 201–203.
Chambers, L.E., Hughes, L. & Weston, M.A. 2005. Climate change and its impact on Australia’s avifauna.
Emu 105, 1–20.
Chan, B.K.K. & Hung, O.S. 2005. Cirral length of the acorn barnacle Tetraclita japonica (Cirripedia: Bala-
nomorpha) in Hong Kong: effect of wave exposure and tidal height. Journal of Crustacean Biology
25, 329–332.
Chavez, F.P., Ryan, J., Lluch-Cota, S.E. & Niquen, M. 2003. From anchovies to sardines and back: multidecadal
change in the Pacific Ocean. Science 299, 217–221.
Choo, K.S., Nilsson, J., Pedersen, M. & Snoeijs, P. 2005. Photosynthesis, carbon uptake and antioxidant
defence in two coexisting filamentous green algae under different stress conditions. Marine Ecology
Church J.A., Gregory, J.M., Huybrechts, P., Kuhn, M., Lambeck, K., Nhuan, M.T., Qin, D. & Woodworth,
P.L. 2001. Changes in sea level. In Climate 2001: The Scientific Basis. Contribution of Working
Group 1 to the Third Assessment Report of the Intergovernmental Panel on Climate Change. J.T.
Houghton et al. (eds) Cambridge, UK & New York, USA: Cambridge University Press, 639-694.
Clarke, P.J., Kerrigan, R.A. & Westphal, C.J. 2001. Dispersal potential and early growth in 14 tropical
mangroves: do early life history traits correlate with patterns of adult distribution? Journal of Ecology
89, 648–659.
Clarke, P.J. & Myerscough, P.J. 1991. Floral biology and reproductive phenology of Avicennia marina in
south-eastern Australia. Australian Journal of Botany 39, 283–293.
Coates, M. 1998. A comparison of intertidal assemblages on exposed and sheltered tropical and temperate
rocky shores. Global Ecology and Biogeography 7, 115–124.
Cole, R.G., Babcock, R.C., Travers, V. & Creese, R.G. 2001. Distributional expansion of Carpophyllum
flexuosum onto wave-exposed reefs in northeastern New Zealand. New Zealand Journal of Marine
Coleman, F.C. & Williams, S.L. 2002. Overexploiting marine ecosystem engineers: potential consequences
for biodiversity. Trends in Ecology and Evolution 17, 40–44.
Cordi, B., Donkin, M.E., Peloquin, J., Price, D.N. & Depledge, M.H. 2001. The influence of UV-B radiation
on the reproductive cells of the intertidal macroalga, Enteromorpha intestinalis. Aquatic Toxicology
56, 1–11.
Costanza, R., d’Arge, R., de Groot, R., Farber, S., Grasso, M., Hannon, B., Limburg, K., Naeem, S., O’Neill,
R.V., Paruelo, J., Raskin, R.G., Sutton, P. & van den Belt, M. 1997. The value of the world’s ecosystem
services and natural capital. Nature 387, 253–260.
Cowen, R.K., Paris, C.B. & Srinivasan, A. 2006. Scaling of connectivity in marine populations. Science 311,
522–527
Crick, H.Q.P., Dudley, C., Glue, D.F. & Thomson, D.L. 1997. U.K. birds are laying eggs earlier. Nature 388, 536.
Cruz-Palacios, V. & Van Tussenbroek, B.I. 2005. Simulation of hurricane-like disturbances on a Caribbean
seagrass bed. Journal of Experimental Marine Biology and Ecology 324, 44–60.
CSIRO. 2006. Wealth from Oceans Flagship overview. Available HTTP: http://www.csiro.au (accessed 10 July
2006).
457
Cubasch, U., Meehl, G.A., Boer, G.J., Stouffer, R.J., Dix, M., Noda, A., Senior, C.A., Raper, S. & Yap, K.S.
2001. Projections of future climate change. In Climate 2001: The Scientific Basis. Contribution of
Working Group 1 to the Third Assessment Report of the Intergovernmental Panel on Climate Change.
J.T. Houghton et al. (eds) Cambridge, UK & New York, USA: Cambridge University Press.
Cushing, D.H. 1990. Plankton production and year-class strength in fish populations: an update of the
match/mismatch hypothesis. Advances in Marine Biology 26, 250–293.
Damkaer, D.M. & Dey, D.B. 1983. UV damage and photoreactivation potentials of larval shrimp, Pandulus
platyceros, and adult euphausiids, Thysanonessa raschii. Oecologia 60, 169–175.
Davenport, J. 1997. Temperature and the life-history strategies of sea turtles. Journal of Thermal Biology 22,
479–488.
Davenport, J. & Davenport, J.L. 2005. Effects of shore height, wave exposure and geographical distance on
thermal niche width of intertidal fauna. Marine Ecology Progress Series, 292, 41–50.
Davis, A.R. & Butler, A.J. 1989. Direct observations of larval dispersal in the colonial ascidian Podoclavella
moluccensis Sluiter: evidence for closed populations. Journal of Experimental Marine Biology and
Ecology 127, 189–203.
Davis, G., Richards, D.V., Haaker, P.L. & Parker, D.O. 1996. Abalone population declines and fishery
management in southern California. In Abalone of the World: Biology, Fisheries and Culture, S.A.
Guzman del Proo (ed.). Cambridge, Massachusetts: Fishing News Books, 237–249.
Dawson, S.P. & Dennison, W.C. 1996. Effects of ultraviolet and photosynthetically active radiation on five
seagrass species. Marine Biology 125, 629–638.
Dayton, P.K. & Tegner, M.J. 1984. Catastrophic storms, El Niño, and patch stability in a southern California
kelp community. Science 224, 283–285.
Dayton, P.K. & Tegner, M.J. 1989. Bottoms beneath troubled waters: Benthic impacts of the 1982–1984 El
Niño in the temperate zone. In Global Ecological Consequences Of the 1982–1983 El-Nino-Southern
Oscillation, P. Glynn (ed.). Amsterdam: Elsevier, 473–472.
Dayton, P.K., Tegner, M.J. & Edwards, P.B. 1999. Temporal and spatial scales of kelp demography: the role
of oceanographic climate. Ecological Monographs 69, 219–250.
Dayton, P.K., Tegner, M.J., Parnell, P.E. & Edwards, P.B. 1992. Temporal and spatial patterns of disturbance
and recovery in a kelp forest community. Ecological Monographs 62, 421–445.
de Jager, D., Manning, M. & Kuijpers, L. 2005. Safeguarding the ozone layer and the global climate system:
issues related to hydrofluorocarbons and perfluorocarbons. IPCC/TEAP Special Report, Technical
Summary.
de Lange, W.P. & de Lange, P.J. 1994. An appraisal of factors controlling the latitudinal distribution of
mangrove (Avicennia marina var. resinifera) in New Zealand. Journal of Coastal Research 10,
539–548.
de Lange, W.P. & Gibb, J.G. 2000. Seasonal, interannual and decadal variability of storm surges at Tauranga,
New Zealand. New Zealand Journal of Marine and Freshwater Research 34, 419–434.
De Vantier, L.M., De’ath, G., Turak, E., Done, T.J. & Fabricius, K.E. 2006. Species richness and community
structure of reef-building corals on the nearshore Great Barrier Reef. Coral Reefs 253, 329–340.
Denny, M. & Gaylord, B. 1996. Why the urchin lost its spines: hydrodynamic forces and survivorship in three
echinoids. Journal of Experimental Biology 199, 717–729.
Detres, Y., Armstrong, R.A. & Connelly, X.M. 2001. Ultraviolet-induced responses in two species of climax
tropical marine macrophytes. Journal of Photochemistry and Photobiology B 62, 55–66.
Devlin, M.J. & Brodie, J. 2005. Terrestrial discharge into the Great Barrier Reef Lagoon: nutrient behaviour
in coastal waters. Marine Pollution Bulletin 51, 9–22.
Deysher, L.E. & Dean, T.A. 1986. Interactive effects of light and temperature on sporophyte production in
the giant kelp Macrocystis pyrifera. Marine Biology 93, 17–20.
Diaz-Almela, E., Marba, N., Alvarez, E., Balestri, E., Ruiz-Fernandez, J.M. & Duarte, C.M. 2006. Patterns of
seagrass (Posidonia oceanica) flowering in the western Mediterranean. Marine Biology 148, 723–742.
Diehl, S. 2002. Phytoplankton, light, and nutrients in a gradient of mixing depths: theory. Ecology 83, 386–391.
Döhler, G. 1994. UV effects on the nitrogen metabolism of marine phytoplankton and adaptation to UV
radiation. In Stratospheric Ozone Depletion/UV-B Radiation in the Biosphere, R. Biggs & M. Joyner
(eds). NATO ASI Series, 18. Berlin: Springer-Verlag, 163–174.
458
Done, T.J., Whetton, P., Jones, R., Berkelmans, R., Lough, J., Skirving, W. & Wooldridge, S. 2003. Global
climate change and coral bleaching on the Great Barrier Reef. Final report to the State of Queensland
Greenhouse Taskforce through the Department of Natural Resources and Mining. AIMS, Townsville,
Australia.
Donner, S.D., Skirving, W.J., Little, C.M., Oppenheimer, M. & Hoegh-Guldberg, O. 2005. Global assessment of
coral bleaching and required rates of adaptation under climate change. Global Change Biology 11, 1–15.
Dorenbosch, M., Grol, M.G.G., Christianen, M.J.A., Nagelkerken, I. & van der Velde, G. 2005. Indo-Pacific
seagrass beds and mangroves contribute to fish density and diversity on adjacent coral reefs. Marine
Drinkwater, K.F. 2005. The response of Atlantic cod (Gadus morhua) to future climate change. ICES Journal
of Marine Science 62, 1327–1337.
Drohan, A.F., Thoney, D.A. & Baker, A.C. 2005. Synergistic effect of high temperature and ultraviolet-B
radiation on the gorgonian Eunicea tourneforti (Octocorallia: Alcyonacea: Plexauridae). Bulletin of
Duarte, C.M., Cebrián, J. and Marbà, N. 1992. Uncertainty of detecting sea change. Nature 356, 190.
Duarte, C.M., Middelburg, J.J. & Caraco, N. 2005. Major role of marine vegetation on the oceanic carbon
cycle. Biogeosciences 1, 173–180.
Duke, N.C. 1992. Mangrove floristics and biogeography. In Tropical Mangrove Ecosystems, A.I. Robertson
& D.M. Alongi (eds). Coastal and Estuarine Studies 41, 63–100.
Duke, N.C., Bell, A.M., Pederson, D.K., Roelfsema, C.M. & Nash, S.B. 2005. Herbicides implicated as the
cause of severe mangrove dieback in the Mackay region, NE Australia: consequences for marine plant
habitats of the GBR World Heritage Area. Marine Pollution Bulletin 51, 308–324.
Dulvy, N.K., Sadovy, Y. & Reynolds, J.D. 2003. Extinction vulnerability in marine populations. Fish and
Fisheries 4, 25–64.
Dunlop, J.N., Surman, C.A. & Wooller, R.D. 2001. The marine distribution of seabirds from Christmas Island,
Indian Ocean. Emu 101, 19–24.
Dunlop, J.N. & Wooller, R.D. 1986. Range extensions and the breeding seasons of seabirds in south-western
Australia. Records of the Western Australian Museum 12, 389–394.
Dunlop, J.N. & Wooller, R.D. 1990. The breeding seabirds of southwestern Australia: trends in species,
populations and colonies. Corella 14, 107–112.
Eckert, S.A. 2002. Distribution of juvenile leatherback sea turtle Dermochelys coriacea sightings. Marine
Edgar, G.J., Moverley, J., Barrett, N.S., Peters, D. & Reed, C. 1997. The conservation-related benefits of a
systematic marine biological sampling programme: the Tasmanian reef bioregionalisation as a case
study. Biological Conservation 79, 227–240.
Edgar, G.J., Samson, C.R. & Barrett, N.S. 2005. Species extinction in the marine environment: Tasmania as
a regional example of overlooked losses in biodiversity. Conservation Biology 19, 1294–1300.
Edwards, M. 2004. Preface. Marine Ecology Progress Series Suppl. 1–2, 1–2.
Edwards, M. & Richardson, A.J. 2004. Impact of climate change on marine pelagic phenology and trophic
mismatch. Nature 430, 881–884.
Edwards, M.J., Johns, D.G., Leterme, S.C., Svendsen, E. & Richardson, A.J. 2006. Regional climate change
and harmful algal blooms in the northeast Atlantic. Limnology and Oceanography 51, 820–829.
Edwards, M.S. 2004. Estimating scale-dependency in disturbance impacts: El Niños and giant kelp forests in
Edyvane, K.S. 2003. Conservation, monitoring and recovery of threatened giant kelp (Macrocystis pyrifera)
beds in Tasmania. Final Report for Environment Australia. Department of Primary Industries, Water
and Environment, Hobart, Tasmania.
El-Sayed, S., van Dijken, G.L. & Gonzalez-Rodas, G. 1996. Effects of increasing ultraviolet radiation on
marine ecosystems. International Journal of Environmental Studies 51, 199–216.
Ellison, A.M. & Farnsworth, E.J. 1993. Seedling survivorship, growth and response to disturbance in Belizean
mangal. American Journal of Botany 80, 1137–1145.
Ellison, A.M. & Farnsworth, E.J. 1997. Simulated sea level change alters anatomy, physiology, growth and
reproduction of red mangrove (Rhizophora mangle L.). Oecologia 112, 435–446.
459
Ellison, J.C. 1993. Mangrove retreat with rising sea level, Bermuda. Estuarine Coastal and Shelf Science 37,
75–87.
Ellison, J.C. & Stoddart, D.R. 1991. Mangrove ecosystem collapse during predicted sea-level rise — holocene
analogs and implications. Journal of Coastal Research 7, 151–165.
Emanuel, K. 2005. Increasing destructiveness of tropical cyclones over the past 30 years. Nature 436, 686–688.
Endean, R. 1973. Destruction and recovery of coral reef communities In Biology and Geology of Coral Reefs,
O.A. Jones & R. Endean (eds). New York: Academic Press, 215–254.
Ene, A., Su, M., Lemaire, S., Rose, C., Schaff, S., Moretti, R., Lenz, J. & Herbst, L.H. 2005. Distribution of
chelonid fibropapillomatosis-associated herpesvirus variants in Florida: molecular genetic evidence
for infection of turtles following recruitment to neritic developmental habitats. Journal of Wildlife
Diseases 41, 489–497.
Engel, A., Zondervan, I., Aerts, K., Beaufort, L., Benthien, A., Chou, L., Delille, B., Gattuso, J.P., Harlay, J.,
Heemann, C., Hoffmann, L., Jacquet, S., Nejstgaard, J., Pizay, M.D., Rochelle-Newall, E., Schneider,
U., Terbrueggen, A. & Riebesell, U. 2005. Testing the direct effect of CO2 concentration on a bloom
of the coccolithophorid Emiliania huxleyi in mesocosm experiments. Limnology and Oceanography
50, 493–507.
Environment Australia. 1998. Draft recovery plan for marine turtles in Australia. Wildlife Management Section,
Biodiversity Group, Environment Australia.
Eppley, R.W. 1972. Temperature and phytoplankton growth in the sea. Fishery Bulletin 70, 1063–1085.
Erga, S.R., Aursland, K., Frette, O., Hamre, B., Lotsberg, J.K., Stamnes, J.J., Aure, J., Rey, F. & Stamnes, K.
2005. UV transmission in Norwegian marine waters: controlling factors and possible effects on
primary production and vertical distribution of phytoplankton. Marine Ecology Progress Series 305,
79–100.
Evans, M.J. & Williams, R.J. 2001. Historical distribution of estuarine wetlands at Kurnell Peninsula, Botany
Bay. Wetlands 19, 61–71.
Ewel, K.C., Twilley, R.R. & Ong, J.E. 1998. Different kinds of mangrove forests provide different goods and
services. Global Ecology and Biogeography 7, 83–94.
Fabricius, K.E. 2005. Effects of terrestrial runoff on the ecology of corals and coral reefs: review and synthesis.
Marine Pollution Bulletin 50, 125–146.
Farnsworth, E.J., Ellison, A.M. & Gong, W.K. 1996. Elevated CO2 alters anatomy, physiology, growth, and
reproduction of red mangrove (Rhizophora mangle L). Oecologia 108, 599–609.
Ferraroli, S., Georges, J.Y., Gaspar, P. & Le Maho, Y. 2004. Endangered species: where leatherback turtles
meet fisheries. Nature 429, 521–522.
Field, C.D. 1995. Impact of expected climate change on mangroves. Hydrobiologia 285, 75–81.
Fish, M.R., Côté, I.M., Gill, J.A., Jones, A.P., Renshoff, S. & Watkinson, A.R. 2005. Predicting the impact
of sea-level rise on Caribbean sea turtle nesting habitat. Conservation Biology 19, 482–491.
Foley, A.M., Schroeder, B.A., Redlow, A.E., Fick-Child, K.J. & Teas, W.G. 2005. Fibropapillomatosis in
stranded green turtles (Chelonia mydas) from the eastern United States (1980–1998): Trends and
associations with environmental factors. Journal of Wildlife Diseases 41, 29–41.
Fonseca, C.R. & Ganade, G. 2001. Species functional redundancy, random extinctions and the stability of
ecosystems. Journal of Ecology 89, 118–125.
Fonseca, M.S. & Bell, S.S. 1998. Influence of physical setting on seagrass landscapes near Beaufort, North
Carolina, U.S.A. Marine Ecology Progress Series 171, 109–121.
Fourqurean, J.W. & Rutten, L.M. 2004. The impact of Hurricane Georges on soft-bottom, back reef commu-
nities: site- and species-specific effects in south Florida seagrass beds. Bulletin of Marine Science 75,
239–257.
Fowler-Walker, M.J., Connell, S.D. & Gillanders, B.M. 2005. To what extent do geographic and associated
environmental variables correlate with kelp morphology across temperate Australia? Marine and
Freshwater Research 56, 877–887.
Fowler-Walker, M.J., Wernberg, T. & Gillanders, B.M. 2006. Differences in kelp morphology between wave
sheltered and exposed localities: morphologically plastic or fixed traits? Marine Biology 148, 755–767.
Frid, C.L.J. & Fordham, E. 1994. The morphology of the sub-littoral gastropod Gibbula cineraria (L.) along
a gradient of wave exposure. Ophelia 40, 135–146.
460
Friedel, M.H., Nelson, D.J., Sparrow, A.D., Kinloch, J.E. & Maconochie, J.R. 1993. What induces central
Australian arid zone trees and shrubs to flower and fruit. Australian Journal of Botany 41, 307–319.
Fulton, E.A., Smith, A.D.M. & Punt, A.E. 2005. Which ecological indicators can robustly detect effects of
fishing? ICES Journal of Marine Science 62, 540–541.
Furnas, M.J. & Crosbie, N.D. 1999. In situ growth dynamics of the photosynthetic prokaryotic picoplankters
Synechococcus and Prochlorococcus. Bulletin de l’Institut Oceanographique Monaco S, 387–417.
Furukawa, K., Wolanski, E. & Mueller, H. 1997. Currents and sediment transport in mangrove forests.
Galbraith, H., Jones, R., Park, R., Clough, J., Herrod-Julius, S., Harrington, B. & Page, G. 2002. Global
climate change and sea level rise: potential losses of intertidal habitat for shorebirds. Waterbirds 25,
173–183.
Gao, K., Argua, Y., Asada, K., Ishihara, T., Akano, T. & Kiyohara, M. 1993. Calcification in the articulated
coralline alga Corallina pilulifera, with special reference to the effect of elevated CO2 concentration.
Gao, K.S., Ji, Y. & Aruga, Y. 1999. Relationship of CO2 concentrations to photosynthesis of intertidal
macroalgae during emersion. Hydrobiologia 399, 355–359.
Garcia, C.M. & de Perera, T.B. 2002. Ultraviolet-based female preferences in a viviparous fish. Behavioral
Ecology and Sociobiology 52, 1–6.
Garde, K. & Cailliau, C. 2000. The impact of UV-B radiation and different PAR intensities on growth, uptake
of 14C, excretion of DOC, cell volume, and pigmentation in the marine prymensiophyte Emiliania
huxleyi. Journal of Experimental Marine Biology and Ecology 247, 99–112.
Gardner, T.A., Côté, I.M., Gill, J.A., Grant, A. & Watkinson, A.R. 2005. Hurricanes and Caribbean coral reefs:
impacts, recovery patterns, and role in long-term decline. Ecology 86, 174–184.
Garnett, S.T. & Crowley, G.M. 2000. The Action Plan for Australian Birds. Canberra, Australia: Environment
Australia.
Garrabou, J., Perez, T., Sartoretto, S. & Harmelin, J.G. 2001. Mass mortality event in red coral Corallium
rubrum populations in the Provence region (France, NW Mediterranean). Marine Ecology Progress
Series 217, 263–272.
Gattuso, J.P., Frankignoulle, M., Bourge, I., Romaine, S. & Buddemeier, R.W. 1998. Effect of calcium
carbonate saturation of seawater on coral calcification. Global and Planetary Change 18, 37–46.
Gaughan, D.J. 2001. Disease-translocation across geographic boundaries must be recognized as a risk even
in the absence of disease identification: the case with Australian Sardinops. Reviews in Fish Biology
and Fisheries 11, 113–123.
Gaylord, B. & Gaines, S.D. 2000. Temperature or transport? Range limits in marine species mediated solely
by flow. American Naturalist 155, 769–789.
Genin, A., Dayton, P.K., Lonsdale, P.F. & Spiess, F.N. 1986. Corals on seamount peaks provide evidence of
current acceleration over deep-sea topography. Nature 322, 59–61.
Gershwin, L.A. 2005. Two new species of jellyfishes (Cnidaria: Cubozoa: Carybdeida) from tropical Western
Australia, presumed to cause Irukandji syndrome. Zootaxa 1084, 1–30.
Gillett, N.P. & Thompson, D.W. 2003. Simulation of recent Southern Hemisphere climate change. Science
302, 273–275.
Gilmour, J. 1999. Experimental investigation into the effects of suspended sediment on fertilisation, larval
survival and settlement in a scleractinian coral. Marine Biology 135, 451–462.
Giordano, M., Norici, A. & Hell, R. 2005. Sulfur and phytoplankton: acquisition metabolism and impact on
the environment. New Phytologist 166, 371–382.
Gleason, D.F., Edmunds, P.J. & Gates, R.D. 2006. Ultraviolet radiation effects on the behavior and recruitment
of larvae from the reef coral Porites astreoides. Marine Biology 148, 503–512.
Glen, F. & Mrosovsky, N. 2004. Antigua revisited: the impact of climate change on sand and nest temperatures
at a hawksbill turtle (Eretmochelys imbricata) nesting beach. Global Change Biology 10, 2036–2045.
Glover, A.G. & Smith, C.R. 2003. The deep-sea floor ecosystem: current status and prospects of anthropogenic
change by the year 2025. Environmental Conservation 30, 219–241.
Godfrey, M.H., Barreto, R. & Mrosovsky, N. 1996. Estimating past and present sex ratios of sea turtles in
Suriname. Canadian Journal of Zoology 74, 267–277.
461
Godfrey, M.H., D’Amato, A.F., Marcovaldi, M.A. & Mrosovsky, N. 1999. Pivotal temperature and predicted
sex ratios for hatchling hawksbill turtles from Brazil. Canadian Journal of Zoology 77, 1465–1473.
Goes, J.I., Handa, N., Taguchi, S. & Hama, T. 1994. Effect of UV-B radiation on the fatty acid composition
of the marine phytoplankton Tetraselmis sp.: relationship to cellular pigments. Marine Ecology
Goffart, A., Hecq, J.H. & Legendre, L. 2002. Changes in the development of the winter-spring phytoplankton
bloom in the Bay of Calvi (NW Mediterranean) over the last two decades: a response to changing
climate? Marine Ecology Progress Series 236, 45–60.
Goggin, C.L. & Lester, R.J.G. 1995. Perkinsus, a protistan parasite of abalone in Australia — a review. Marine
Goldberg, N.A. & Kendrick, G.A. 2004. Effects of island groups, depth, and exposure to ocean waves on
subtidal macroalgal assemblages in the Recherche Archipelago, Western Australia. Journal of Phy-
cology 40, 631–641.
Gordon, H.B., Rotstayn, L.D., McGregor, J.L., Dix, M.R., Kowalczyk, E.A., O’Farrell, S.P., Waterman, L.J.,
Hirst, A.C., Wilson, S.G., Collier, M.A., Watterson, I.G. & Elliott, T.I. 2002. The CSIRO Mk3 Climate
System Model. CSIRO Atmospheric Research, Australia, Technical Paper No. 60.
Gorgula, S.K. & Connell, S.D. 2004. Expansive covers of turf-forming algae on human-dominated coast: the
relative effects of increasing nutrient and sediment loads. Marine Biology 145, 613–619.
Graham, M.H. 1996. Effect of high irradiance on recruitment of the giant kelp Macrocystis (Phaeophyta) in
shallow water. Journal of Phycology 32, 903–906.
Graham, M.H. 1997. Factors determining the upper limit of giant kelp, Macrocystis pyrifera Agardh, along
the Monterey Peninsula, central California, U.S.A. Journal of Experimental Marine Biology and
Ecology 218, 127–149.
Gregory, J.M., Church, J.A., Boer, G.J., Dixon, K.W., Flato, G.M., Jackett, D.R., Lowe, J.A., O’Farrell, S.P.,
Roeckner, E., Russell, G.L., Stouffer, R.J. & Winton, M. 2001. Comparison of results from several
AOGCMs for global and regional sea-level change 1900–2100. Climate Dynamics 18, 223–240.
Greve, W., Reiners, F., Nast, J. & Hoffmann, S. 2004. Helgoland Roads meso- and macrozooplankton time-
series 1974 to 2004: lessons from 30 years of single spot, high frequency sampling at the only off-
shore island of the North Sea. Helgoland Marine Research 58, 274–288.
Griffies, S.M., Gnanadesikan, A., Pacanowski, R.C., Larichev, V.D., Dukowicz, J.K. & Smith, R.D. 1998.
Isoneutral diffusion in a z-coordinate ocean model. Journal of Physical Oceanography 28, 805–830.
Griffin, D.A., Wilkin, J.L., Chubb, C.F., Pearce, A.F. & Caputi, N. 2001. Ocean currents and the larval phase
of Australian rock lobster, Panulirus cygnus. Marine and Freshwater Research 52, 1187–1199.
Griffiths, S.P. 2003. Rockpool ichthyofaunas of temperate Australia: species composition, residency and
biogeographic patterns. Estuarine, Coastal and Shelf Science 58, 173–186.
Grove, R.S., Zabloudil, K., Norall, T. & Deysher, L. 2002. Effects of El Niño events on natural kelp beds and
artificial reefs in southern California. ICES Journal of Marine Science 59, S330–S337.
Grover, J. 1997. Resource Competition. London: Chapman & Hall.
Guinotte, J.M., Orr, J., Cairns, S., Freiwald, A., Morgan, L. & George, R. 2006. Will human-induced changes
in seawater chemistry alter the distribution of deep-sea scleractinian corals? Frontiers in Ecology and
the Environment 4, 141–146.
Häder, D. & Häder, M.A. 1989. Effects of solar UV-B irradiation on photomovement and motility in photo-
synthetic and colorless flagellates. Environmental and Experimental Botany 29, 273–282.
Häder, D. & Liu, S.L. 1990. Effects of artificial and solar UV-B radiation on the gravitactic orientation of the
dinoflagellate Peridinium gatunense. FEMS Microbiology, Ecology 73, 331–338.
Häder, D.P., Kumar, H.D., Smith, R.C. & Worrest, R.C. 1998. Effects on aquatic ecosystems. Journal of
Photochemistry and Photobiology B 46, 53–68.
Hall, S.J. 2002. The continental shelf benthic ecosystem: current status, agents for change and future prospects.
Environmental Conservation 29, 350–374.
Hanelt, D., Wiencke, C. & Nultsch, W. 1997. Influence of UV radiation on the photosynthesis of Arctic
macroalgae in the field. Journal of Photochemistry and Photobiology B 38, 40–47.
Hansen, J., Sato, M., Ruedy, R., Lo, K., Lea, D.W. & Medina-Elizade, M. 2006. Global temperature change.
Proceedings of the National Academy of Sciences of the United States of America 103, 14,288–14,293.
462
Harborne, A.R., Mumby, P.J., Micheli, F., Perry, C.T., Dahlgren, C.P., Holmes, K.E. & Brumbaugh, D.R.
2006. The functional value of Caribbean coral reef, seagrass and mangrove habitats to ecosystem
processes. Advances in Marine Biology 50, 57–189.
Harris, G., Nilsson, C., Clementson, L. & Thomas, D. 1987. The water masses of the east-coast of Tasmania:
seasonal and interannual variability and the influence on phytoplankton biomass and productivity.
Australian Journal of Marine and Freshwater Research 38, 569–590.
Harris, G.P., Davies, P., Nunez, M. & Meyers, G. 1988. Interannual variability in climate and fisheries in
Tasmania. Nature 333, 754–757
Harris, G.P., Griffiths, F.B. & Clementson, L.A. 1992. Climate and the fisheries off Tasmania: interactions of
physics, food chains and fish. South African Journal of Marine Science 12, 585–597.
Harris, G.P., Griffiths, F.B., Clementson, L.A., Lyne, V. & Vanderhoe, H. 1991. Seasonal and interannual
variability in physical processes, nutrient cycling and the structure of the food chain in Tasmanian
shelf waters. Journal of Plankton Research 13(Suppl.), S109–S131.
Harty, C. 2004. Planning strategies for mangrove and saltmarsh changes in southeast Australia. Coastal
Management 32, 405–415.
Harty, C. & Cheng, D. 2003. Ecological assessment and strategies for the management of mangroves in
Brisbane Water — Gosford, New South Wales, Australia. Landscape and Urban Planning 62, 219–240.
Harvell, C.D., Kim, K., Burkholder, J.M., Colwell, R.R., Epstein, P.R., Grimes, D.J., Hofmann, E.E., Lipp,
E.K., Osterhaus, A.D.M.E., Overstreet, R.M., Porter, J.W., Smith, G.W. & Vasta, G.R. 1999. Review:
marine ecology — emerging marine diseases — climate links and anthropogenic factors. Science 285,
1505–1510.
Harvell, C.D., Mitchell, C.E., Ward, J.R., Altizer, S., Dobson, A.P., Ostfeld, R.S. & Samuel, M.D. 2002.
Climate warming and disease risks for terrestrial and marine biota. Science 296, 2158–2162.
Hays, G.C. 2000. The implications of variable remigration intervals for the assessment of population size in
marine turtles. Journal of Theoretical Biology 206, 221–227.
Hays, G.C., Broderick, A.C., Glen, F. & Godley, B.J. 2003. Climate change and sea turtles: a 150-year
reconstruction of incubation temperatures at a major marine turtle rookery. Global Change Biology
9, 642–646.
Hays, G.C., Broderick, A.C., Glen, F., Godley, B.J., Houghton, J.D. & Metcalfe, J.D. 2002. Water temperature
and internesting intervals for loggerhead (Caretta caretta) and green (Chelonia mydas) sea turtles.
Journal of Thermal Biology 27, 429–432.
Hays, G.C., Richardson, A.J. & Robinson, C. 2005. Climate change and marine plankton. Trends in Ecology
and Evolution 20, 337–344.
Hayward, T. 1997. Pacific Ocean climate change: atmospheric forcing, ocean circulation and ecosystem
response. Trends in Ecology and Evolution 12, 150–154.
Helmuth, B., Kingsolver, J.G. & Carrington, E. 2005. Biophysics, physiological ecology, and climate change:
does mechanism matter? Annual Review of Physiology 67, 177–201.
Henry, G.W. & Lyle, J.M. (eds) 2003. National Recreational Fishing Survey. In The National Recreational
and Indigenous Fishing Survey. Canberra: Australian Government Department of Agriculture, Fish-
eries and Forestry, FRDC Project No. 99/158, 27–97.
Herbst, L., Ene, A., Su, M., Desalle, R. & Lenz, J. 2004. Tumor outbreaks in marine turtles are not due to
recent herpesvirus mutations. Current Biology 14, R697–R699.
Hernando, M.P. & Ferreyra, G.A. 2005. The effects of UV radiation on photosynthesis in an Antarctic diatom
(Thalassiosira sp.): does vertical mixing matter? Journal of Experimental Marine Biology and Ecology
325, 35–45.
Hessen, D., de Lange, H.J. & van Donk, E. 1997. UV-induced changes in phytoplankton cells and its effects
on grazers. Freshwater Biology 38, 513–524.
Hewavisenthi, S. & Parmenter, C.J. 2002a. Thermosensitive period for sexual differentiation of the gonads of
the flatback turtle (Natator depressus Garman). Australian Journal of Zoology 50, 521–527.
Hewavisenthi, S. & Parmenter, C.J. 2002b. Incubation environment and nest success of the flatback turtle
(Natator depressus) from a natural nesting beach. Copeia 2002, 302–312.
Hickling, R., Roy, D.B., Hill, J.K., Fox, R. & Thomas, C.D. 2006. The distributions of a wide range of
taxonomic groups are expanding polewards. Global Change Biology 12, 450–455.
463
Hinga, K.R. 2002. Effects of pH on coastal marine phytoplankton. Marine Ecology Progress Series 238,
281–300.
Hirst, A.G. & Lampitt, R.S. 1998. Towards a global model of in situ weight-specific growth in marine
planktonic copepods. Marine Biology 132, 247–257.
Hiscock, K., Southward, A., Tittley, I. & Hawkins, S. 2004 Effects of changing temperature on benthic marine
life in Britain and Ireland. Aquatic Conservation Marine and Freshwater Ecosystems 14, 333–362.
Hobday, A.J. & Hartmann, K. 2006. Near real-time spatial management based on habitat predictions for a
longline bycatch species. Fisheries Management and Ecology 13, 365–380.
Hobday, A.J., Okey, T.A., Poloczanska, E.S., Kunz, T.J. & Richardson, A.J. (eds) 2006. Impacts of climate
change on Australian marine life. Report to the Australian Greenhouse Office, Canberra, Australia.
Hoegh-Guldberg, O. 1999. Coral bleaching, climate change and the future of the world’s coral reefs. Marine
Hoegh-Guldberg, O. 2004. Marine ecosystems and climate change. In Climate Change and Biodiversity, T.E.
Lovejoy & L. Hannah (eds). New Haven, Connecticut: Yale University Press.
Hoegh-Guldberg, H. & Hoegh-Guldberg, O. 2004. Biological, Economic and Social Impacts of Climate
Change on the Great Barrier Reef. Sydney, Australia: World Wide Fund for Nature.
Hogue, V.E., Wilkerson, F.P. & Dugdale, R.C. 2005. Ultraviolet-B radiation effects on natural phytoplankton
assemblages of central San Francisco Bay. Estuaries 28, 190–203.
Holbrook, S.J., Schmitt, R.J. & Stephens, Jr., J.S. 1997. Changes in an assemblage of temperature reef fishes
associated with a climate shift. Ecological Applications 7, 1299–1310.
Huesemann, M.H., Skillman, A.D. & Crecelius, E.A. 2002. The inhibition of marine nitrification by ocean
disposal of carbon dioxide. Marine Pollution Bulletin 44, 142–148.
Hughes, L. 2003. Climate change and Australia: trends, projections and impacts. Austral Ecology 28, 423–443.
Hughes, T.P., Baird, A.H., Bellwood, D.R., Card, M., Connolly, S.R., Folke, C., Grosberg, R., Hoegh-Guldberg,
O., Jackson, J.B.C., Kleypas, J., Lough, J.M., Marshall, P., Nystrom, M., Palumbi, S.R., Pandolfi,
J.M., Rosen, B. & Roughgarden, J. 2003. Climate change, human impacts, and the resilience of coral
reefs. Science 5635, 929–933.
Hughes, T.P. & Connell, J.H. 1999. Multiple stressors on coral reefs: a long-term perspective. Limnology and
Huisman, J. & Weissing, F.J. 1995. Competition for nutrients and light in a mixed water column: a theoretical
analysis. American Naturalist 146, 536–564.
Hunter, J.R., Kaupp, S.E. & Taylor, J.H. 1982. Assessment of effects of UV radiation on marine fish larvae.
In The Role of Solar Ultraviolet Radiation in Marine Ecosystems, J. Calkins (ed.). New York: Plenum,
459–493.
Huntley, M. & Lopez, M.D.G. 1992. Temperature-dependent production of marine copepods: a global syn-
thesis. American Naturalist 140, 201–242.
Inglis, G.J. & Smith, M.P.L. 1998. Synchronous flowering of estuarine seagrass meadows. Aquatic Botany
60, 37–48.
Invers, O., Romero, J. & Pérez, M. 1997. Effects of pH on seagrass photosynthesis, a laboratory and field
assessment. Aquatic Botany 59, 185–194.
Invers, O., Tomas, F., Pérez, M. & Romero, J. 2002. Potential effect of increased global CO2 availability on
the depth distribution of the seagrass Posidonia oceania (L.) Delile, a tentative assessment using a
carbon balance model. Bulletin of Marine Science 71, 1191–1198.
IPCC 2001. Climate 2001: Synthesis Report. A Contribution of Working Groups I, II, and III to the Third
Assessment Report of the Intergovernmental Panel on Climate Change, R.T. Watson & the Core
Writing Team (eds) Cambridge, UK & New York, USA: Cambridge University Press, 398 pp.
IPPC, 2007 Climate Change 2007: The Physical Science Basis: Summary for Policymakers. Contribution of
Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change.
Geneva Switzerland: IPPC, 21 pp.
Jenkins, G.P. 2005. Influence of climate on the fishery recruitment of a temperate, seagrass-associated fish,
the King George whiting Sillaginodes punctata. Marine Ecology Progress Series 288, 263–271.
Jerlov, N.G. & Steeman-Nielsen, E. (eds) 1974. Optical Aspects of Oceanography. New York: Academic Press.
464
Jernakoff, P., Brearley, A. & Nielsen, J. 1996. Factors affecting grazer-epiphyte interactions in temperate
seagrass meadows. Oceanography and Marine Biology: An Annual Review 34, 109–162.
Jickells, T.D., An, Z.S., Andersen, K.K., Baker, A.R., Bergametti, G., Brooks, N., Cao, J.J., Boyd, P.W., Duce,
R.A., Hunter, K.A., Kawahata, H., Kubilay, N., laRoche, J., Liss, P.S., Mahowald, N., Prospero, J.M.,
Ridgwell, A.J., Tegen, I. & Torres, R. 2005. Global iron connections between desert dust, ocean
biogeochemistry, and climate. Science 308, 67–71.
Johnson, C., Ling, S., Ross, J., Shepherd, S. & Miller, K. 2005. Establishment of the long-spined sea urchin
(Centrostephanus rodgersii) in Tasmania: first assessment of potential threats to fisheries. Tasmanian
Aquaculture and Fisheries Institute, Australia, FRDC Project No. 2001/044.
Jones, A.G. 2004. Sea turtles: old viruses and new tricks. Current Biology 14, R842–R843.
Jones, J.M. & Widmann, M. 2004. Early peak in Antarctic oscillation index. Nature 432, 290–291.
Jones, R.J., Bowyer, J., Hoegh-Guldberg, O. & Blackall, L.L. 2004. Dynamics of a temperature-related coral
disease outbreak. Marine Ecology Progress Series 281, 63–77.
Jonsson, P.R., Granhag, L., Moschella, P.S., Aberg, P., Hawkins, S.J. & Thompson, R.C. 2006. Interactions
between wave action and grazing control the distribution of intertidal macroalgae. Ecology 87,
1169–1178.
Jonzén, N., Lindén, A., Torbjørn, E., Knudsen, E., Vik, J.O., Rubolini, D., Piacentini, D., Brinch, C., Spina,
F., Karlsson, L., Stervander, M., Andersson, A., Waldenström, J., Lehikoinen, A., Edvardsen, E.,
Solvang, R. & Stenseth, N.C. 2006. Rapid advance of spring arrival dates in long-distance migratory
birds. Science 312, 1959–1961.
Karanas, J.J., Van Dyke, H. & Worrest, R.C. 1979. Midultraviolet (UV-B) sensitivity of Acartia clausii
Giesbrecht (Copepoda). Limnology and Oceanography 24, 1104–1116.
Karanas, J.J., Worrest, R.C. & Van Dyke, H. 1981. Impact of UV-B radiation on the fecundity of the copepod
Acartia clausii. Marine Biology 65, 125–133.
Karoly, D.J. & Stott, P.A. 2006. Anthropogenic warming of central England temperature. Atmospheric Science
Letters 7, 81–85.
Kathiresan, K. & Bingham, B.L. 2001. Biology of mangroves and mangrove ecosystems. Advances in Marine
Biology 40, 81–251.
Kathiresan, K. & Rajendran, N. 2005. Coastal mangrove forests mitigated tsunami. Estuarine, Coastal and
Kathiresan, K., Rajendran, N. & Thangaduri, G. 1996. Growth of mangrove seedlings in the intertidal area
of Vellar estuary, southeast coast of India. Indian Journal of Marine Sciences 25, 240–243.
Keatley, M.R., Fletcher, T.D., Hudson, I.L. & Ades, P.K. 2002. Phenological studies in Australia: potential
application in historical and future climate analysis. International Journal of Climatology 22,
1769–1780.
Kelfkens, G., Bregman, A., de Gruijl, F.R., van der Leun, J.C., Piquet, A., van Oijen, T., Gieskes, W.W.C.,
van Loveren, H., Velders, G.J.M., Martens, P. & Slaper, H. 2002. Ozone layer — climate change
interactions. Influence on UV levels and UV related effects. Netherlands Environmental Assessment
Agency, Report No. 410200112.
Keller, A.A., Hargraves, P., Jeon, H., Klein-MacPhee, G., Klos, E., Oviatt, C. & Zhang, J. 1997. Effects of
ultraviolet-B enhancement on marine trophic levels in a stratified coastal system. Marine Biology 130,
277–287.
Kemp, D.J. 2001. Reproductive seasonality in the tropical butterfly Hypolimnas bolina (Lepidoptera: Nymph-
alidae) in northern Australia. Journal of Tropical Ecology 17, 483–494.
Kendall, M.A., Burrows, M.T., Southward, A.J. & Hawkins, S.J. 2004. Predicting the effects of marine climate
change on the invertebrate prey of the birds of rocky shores. Ibis 146, 40–47.
Kennelly, S.J. 1987a. Physical disturbances in an Australian kelp community 1. Temporal effects. Marine
Kennelly, S.J. 1987b. Physical disturbances in an Australian kelp community 2. Effects on understorey species
due to differences in kelp cover. Marine Ecology Progress Series 40, 155–165.
Kennelly, S.J. 1989. Effects of kelp canopies on understorey species due to shade and scour. Marine Ecology
465
Keough, M.J. & Butler, A.J. 1995. Temperate subtidal hard substrata. In State of the Marine Environment
Report for Australia: The Marine Environment — Technical Annex: 1, L.P. Zann & P. Kailola (eds).
Canberra: Great Barrier Reef Marine Park Authority for the Department of Environment, Sport and
Territories, Ocean Rescue 2000 Program.
Keser, M., Swenarton, J.T. & Foertch, J.F. 2005. Effects of thermal input and climate change on growth of
Ascophyllum nodosum (Fucales, Phaeophyceae) in eastern Long Island Sound (U.S.A.). Journal of
Sea Research 54, 211–220.
Kester, D.R. 1986. Equilibrium models in seawater: applications and limitations. In The Importance of
Chemical ‘Speciation’ in Environmental Processes, M. Bernhard (ed.). Berlin: Springer Verlag, 337–363.
Kikkawa, T., Ishimatsu, A. & Kita, J. 2003. Acute CO2 tolerance during the early developmental stages of
four marine teleosts. Environmental Toxicology 18, 375–382.
King, B.R., Hicks, J.T. & Cornelius, J. 1992. Population changes, breeding cycles and breeding success over
six years in a seabird colony at Michaelmas Bay, Queensland. Emu 92, 1–10.
Kirby, R.R., Beaugrand, G., Lindley, J.A., Richardson, A.J., Edwards, M. & Reid, P.C. 2007. Climate effects
and benthic-pelagic coupling in the North Sea. Marine Ecology Progress Series 330, 31–38.
Kirkman, H. 1997. Seagrasses of Australia. State of the Environment Technical Paper Series (Estuaries and
the Sea). Canberra, Australia: Department of the Environment.
Kleppel, G.S., Davis, C.S. & Carter, K. 1996. Temperature and copepod growth in the sea: a comment on the
temperature-dependent model of Huntley and Lopez. American Naturalist 148, 397–406.
Kleypas, J.A., Buddemeier, R.W., Archer, D., Gattuso, J.P., Langdon, C. & Opdyke, B.N. 1999. Geochemical
consequences of increased atmospheric carbon dioxide on coral reefs. Science 284, 118–120.
Kleypas, J.A., Buddemeier, R.W., Eakin, C.M., Gattuso, J.-P., Guinotte, J., Hoegh-Guldberg, O., Iglesias-
Prieto, R., Jokiel, P.L., Langdon, C., Skirving, W. & Strong, A.E. 2005. Comment on “Coral reef
calcification and climate change: the effect of ocean warming”. Geophysical Research Letters 32, 1–3.
Knowlton, N. 2001. The future of coral reefs. Proceedings of the National Academy of Sciences of the United
States of America 98, 5419–5425.
Koslow, J.A. 1997. Seamounts and the ecology of deep-sea fisheries. American Scientist 85, 168–176.
Koslow, J.A., Gowlett-Holmes, K., Lowry, J.K., O’Hara, T., Poore, G.C.B. & Williams, A. 2001. Seamount
benthic macrofauna off southern Tasmania: community structure and impacts of trawling. Marine
Koslow, J.A., Hobday, A.J. & Boehlert, G.W. 2002. Climate variability and marine survival of coho salmon
(Oncoryhnchus kisutch) in the Oregon Production Area. Fisheries Oceanography 11, 65–77.
Kouwenberg, J.H.M., Browman, H.I., Cullen, J.J., Davis, R.F., St.-Pierre, J.F. & Runge, J.A. 1999a. Biological
weighting of ultraviolet (280–400 nm) induced mortality in marine zooplankton and fish. I. Atlantic
cod (Gadus morhua) eggs. Marine Biology 134, 269–284.
Kouwenberg, J.H.M., Browman, H.I., Runge, J.A., Cullen, J.J., Davis, R.F. & St.-Pierre, J.F. 1999b. Biological
weighting of ultraviolet (280–400 nm) induced mortality in marine zooplankton and fish. II. Calanus
finmarchicus (Copepoda) eggs. Marine Biology 134, 285–293.
Kuffner, I.B. 2001. Effects of ultraviolet (UV) radiation on larval settlement of the reef coral Pocillopora
damicornis. Marine Ecology Progress Series 217, 251–261.
Kunz, T.J. 2005. Effects of mixing depth, turbulent diffusion and nutrient enrichment on enclosed marine
plankton communities. Ph.D. thesis, Ludwig-Maximilians University, Munich, Germany.
Kurihara, H., Shimode, S. & Shirayama, Y. 2004. Sub-lethal effects of elevated concentration of CO2 on
planktonic copepods and sea urchins. Journal of Oceanography 60, 743–750.
Langdon, C., Takahashi, T., Sweeney, C., Chipman, D., Goddard, J., Marubini, F., Aceves, H., Barnett, H. &
Atkinson, M.J. 2000. Effect of calcium carbonate saturation state on the calcification rate of an
experimental coral reef. Global Biogeochemical Cycles 14, 639–654.
Larkum, A.W.D. 1976. Ecology of Botany Bay I. Growth of Posidonia australis (Brown) Hook f. in Botany Bay
and other Bays of the Sydney Basin. Australian Journal of Marine and Freshwater Research 27, 117–127.
Learmonth, J.A., Macleod, C.D., Santos, M.B., Pierce, G.J., Crick, H.Q.P. & Robinson, R.A. 2006. Potential
effects of climate change on marine mammals. Oceanography and Marine Biology: An Annual Review
44, 431–464.
466
Lee, K.K., Liu, P.C., Chen, Y.C. & Huang, C.Y. 2001. The implication of ambient temperature with the
outbreak of vibriosis in cultured small abalone Haliotis diversicolor supertexta Lischke. Journal of
Thermal Biology 26, 585–587.
Lehikoinen, E., Sparks, T.H. & Zalakevicius, M. 2004. Arrival and departure dates. Birds and Climate Change
35, 1–31.
Lehodey, P. 2001. The pelagic ecosystem of the tropical Pacific Ocean: dynamic and spatial modelling and
biological consequences of ENSO. Progress in Oceanography 49, 439–468.
Lehodey, P., Bertignac, M., Hampton, J., Lewis, A. & Picaut, J. 1997. El Nino Southern Oscillation and tuna
in the western Pacific. Nature 389, 715–718.
Lesser, M.P. 1996. Elevated temperatures and ultraviolet radiation cause oxidative stress and inhibit photo-
synthesis in symbiotic dinoflagellates. Limnology and Oceanography 41, 271–283.
Lesser, M.P. 1997. Oxidative stress causes coral bleaching during exposure to elevated temperatures. Coral
Reefs 16, 187–192.
Lesser, M.P., Kruse, V.A. & Barry, T.M. 2003. Exposure to ultraviolet radiation causes apoptosis in developing
sea urchin embryos. Journal of Experimental Biology 206, 4097–4103.
Li, N.K. & Denny, M.W. 2004. Limits to phenotypic plasticity: flow effects on barnacle feeding appendages.
Biological Bulletin 206, 121–124.
Limpus, C.J. 1992. The hawksbill turtle, Eretmochelys imbricata, in Queensland — population structure within
a southern Great Barrier Reef feeding ground. Wildlife Research 19, 489–506.
Limpus, C.J. & Nicholls, N. 1988. The Southern Oscillation regulates the annual numbers of green turtles
(Chelonia mydas) breeding around northern Australia. Australian Journal of Wildlife Research 15,
157–161.
Limpus, C.J. & Reed, P.C. 1985. Green turtles stranded by cyclone Kathy on the south-western coast of the
Gulf of Carpentaria. Australian Wildlife Research 12, 523–533.
Linderholm, H.W. 2006. Growing season changes in the last century. Agricultural and Forest Meteorology
137, 1–14.
Litchman, E. & Neale, P.J. 2005. UV effects on photosynthesis and acclimation of an estuarine diatom and
cryptomonad. Marine Ecology Progress Series 300, 53–62.
Little, M., Pereira, P., Carrette, T. & Seymour, J. 2006. Jellyfish responsible for Irukandji syndrome. QJM-AN
International Journal of Medicine 99, 425–427.
Logan, B.W. 1961. Cryptozoon and associate stromalolites from the recent, Shark Bay, Western Australia.
Journal of Geology 69, 517–533.
Lohrer, A.M., Thrush, S.F., Hewitt, J.E., Berkenbusch, K., Ahrens, M. & Cummings, V.J. 2004. Terrestrially
derived sediment: response of marine macrobenthic communities to thin terrigenous deposits. Marine
Loneragan, N.R., Adnan, N.A., Connolly, R.M. & Manson, F.J. 2005. Prawn landings and their relationship
with the extent of mangroves and shallow waters in western peninsular Malaysia. Estuarine, Coastal
and Shelf Science 63, 187–200.
Loop, K.A., Miller, J.D. & Limpus, C.J. 1995. Nesting by the Hawksbill Turtle (Eretmochelys imbricata) on
Milman Island, Great Barrier Reef, Australia. Wildlife Research 22, 241–252.
Losey, G.S. 2003. Crypsis and communication functions of UV-visible coloration in two coral reef damselfish,
Dascyllus aruanus and D reticulatus. Animal Behaviour 66, 299–307.
Losey, G.S., Cronin, T.W., Goldsmith, T.H., Hyde, D., Marshall, N.J. & McFarland, W.N. 1999. The UV
visual world of fishes: a review. Journal of Fish Biology 54, 921–943.
Losey, G.S., McFarland, W.N., Loew, E.R., Zamzow, J.P., Nelson, P.A. & Marshall, N.J. 2003. Visual biology
of Hawaiian coral reef fishes. I. Ocular transmission and visual pigments. Copeia Sept. 5, 433–454.
Lough, J.M. 2000. 1997–1998: unprecedented thermal stress to coral reefs? Geophysical Research Letters 27,
3901–3904.
Lough, J.M. & Barnes, D.J. 2000. Environmental controls on growth of the massive coral Porites. Journal of
Luschi, P., Hays, G.C. & Papi, F. 2003. A review of long-distance movements by marine turtles, and the
possible role of ocean currents. Oikos 103, 293–302.
467
Lyle, J.M., Henry, G.W., West, L.D., Campbell, D., Reid, D.D. & Murphy, J.J. 2003.National Recreational
Fishing Survey, In The National Recreational and Indigenous Fishing Survey. G.W. Henry & J.M.
Lyle (eds). Canberra, Australia: Australian Government Department of Agriculture, Fisheries and
Forestry, FRDC Project 99/158, 27-97.
MacLeod, C.D., Bannon, S.M., Pierce, G.J., Schweder, C., Learmonth, J.A., Herman, J.S. & Reid, R.J. 2005.
Climate change and the cetacean community of north-west Scotland. Biological Conservation 124,
477–483.
Majkowski, J., Williams, K. & Murphy, G.I. 1981. Research identifies changing patterns in Australian tuna
fishery. Australian Fisheries 40, 5–10.
Manson, F.J., Loneragan, N.R., Harch, B.D., Skilleter, G.A. & Williams, L. 2005. A broad-scale analysis of
links between coastal fisheries production and mangrove extent: a case-study for northeastern Aus-
tralia. Fisheries Research 74, 69–85.
Mantua, N.J., Hare, S.R., Zhang, Y., Wallace, J.M. & Francis, R.C., 1997. A Pacific interdecadal climate oscillation
with impacts on salmon production. Bulletin American Meteorological Society, 78, 1069–1079.
Marchinko, K.B. & Palmer, A.R. 2003. Feeding in flow extremes: dependence of cirrus form on wave-exposure
in four barnacle species. Zoology 106, 127–141.
Markkula, S.E., Salo, H.M., Immonen, A.K. & Jokinen, E.M. 2005. Effects of short- and long-term ultraviolet
B irradiation on the immune system of the common carp (Cyprinus carpio). Photochemistry and
Photobiology 81, 595–602.
Marra, P.P., Francis, C.M., Mulvihill, R.S. & Moore, F.R. 2005. The influence of climate on the timing and
rate of spring bird migration. Oecologia 142, 307–315.
Marsh, H., Corkeron, P.J., Limpus, C.J., Shaughnessy, P.D. & Ward, T.M. 2001. The reptiles and mammals
in Australian Seas: their status and management. In The State of the Marine Environment Report for
Australia Technical Annex: 1 The Marine Environment, L.P. Zann & P. Kailola (eds). Department of
the Environment, Sport and Territories, Ocean Rescue 2000 Program. Townsville, Queensland, Aus-
tralia: Great Barrier Reef Marine Park Authority, 151–166.
Mason, C.F. 1995. Long-term trends in the arrival dates of spring migrants. Bird Study 42, 182–189.
Maxwell, J.G.H. & Cresswell, G.R. 1981. Dispersal of tropical fauna to the Great Australian Bight by the
Leeuwin Current. Australian Journal of Marine and Freshwater Research 32, 493–500.
McArthur, L.C. & Boland, J.W. 2006. The economic contribution of seagrass to secondary production in South
Australia. Ecological Modelling 196, 163–172.
McGowan, J.A., Cayan, D.R. & Dorman, L.M. 1998. Climate-ocean variability and ecosystem response in
North Pacific. Science, 281, 201–217.
McKenzie, R.L., Björn, L.O., Bais, A. & Ilyasd, M. 2003. Changes in biologically active ultraviolet radiation
reaching the Earth’s surface. Photochemical and Photobiological Sciences 2, 5–15.
McKinnon, A.D. 1996. Growth and development in the subtropical copepod Acrocalanus gibber. Limnology
McKinnon, A.D. & Ayukai, T. 1996. Copepod egg production and food resources in Exmouth Gulf, Western
Australia. Marine and Freshwater Research 47, 595–603.
McKinnon, A.D. & Duggan, S. 2001. Summer egg production rates of paracalanid copepods in subtropical
waters adjacent to Australia’s North West Cape. Hydrobiologia 453/454, 121–132.
McKinnon, A.D. & Duggan, S. 2003. Summer copepod production in subtropical waters adjacent to Australia’s
North West Cape. Marine Biology 143, 897–907.
McKinnon, A.D., Duggan, S. & De’ath, G. 2005. Mesozooplankton dynamics in inshore waters of the Great
Barrier Reef. Estuarine, Coastal and Shelf Science 63, 497–511.
McKinnon, A.D., Richardson, A.J., Burford, M.A. & Furnas, M.J. (in press). Plankton. In Assessing Climate
Change Vulnerability of the Great Barrier Reef. P. Marshall & J. Johnson (eds). Townsville, Queen-
sland, Australia: Great Barrier Reef Marine Park Authority.
McKinnon, A.D. & Thorrold, S.R. 1993. Zooplankton community structure and copepod egg production in
coastal waters of the central Great Barrier Reef lagoon. Journal of Plankton Research 15, 1387–1411.
McMahon, C.R. & Hays, G.C. 2006. Thermal niche, large-scale movements and implications of climate change
for a critically endangered marine vertebrate. Global Change Biology 12, 1–9.
468
McMillan, C. 1984. The distribution of tropical seagrasses with relation to their tolerance of high temperatures.
Aquatic Botany 19, 369–380.
McNeil, B.I. & Matear, R.J. 2006. Projected climate change impact on oceanic acidification. Carbon Balance
and Management, 1, 1–6.
McNeil, B.I., Matear, R.J. & Barnes, D.J. 2004. Coral reef calcification and climate change: the effect of
ocean warming. Geophysical Research Letters 31, L22309.
McNeil, B.I., Matear, R.J., Key, R.M., Bullister, J.L. & Sarmiento, J.L. 2003. Anthropogenic CO2 uptake by
the ocean based on the global chlorofluorocarbon data set. Science 299, 235–239.
McQuoid, M. 2005. Influence of salinity on seasonal germination of resting stages and composition of
microplankton on the Swedish west coast. Marine Ecology Progress Series 289, 151–163.
McShane, P.E., Black, K.P. & Smith, M.G. 1988. Recruitment processes in Haliotis rubra (Mollusca: Gas-
tropoda) and regional hydrodynamics in southeastern Australia imply localised dispersal of larvae.
Menzel, A., Sparks, T.H., Estrella, N., Koch, E., Aasa, A., Ahas, R., Alm-Kübler, K., Bissolli, P., Braslavska, O.,
Briede, A., Chmielewski, F.M., Crepinsek, Z., Curnel, Y., Dalh, A., Defile, C., Donnelly, A., Filella, Y.,
Jatczak, K., Måge, F., Mestre, A., Nordli, Ø., Pe–uelas, J., Pirinen, P., Remis̆ová, V., Scheifinger, H.,
Striz, M., Susnik, A., van Vliet, A.J.H., Wielgolaski, F.-E., Zach, S. & Zust, A. 2006. European pheon-
ological response to climate change matches the warming pattern. Global Change Biology 12, 1–8.
Michaelidis, B., Ouzounis, C., Paleras, A. & Pörtner, H.O. 2005. Effects of long-term moderate hypercapnia
on acid-base balance and growth rate in marine mussels Mytilus galloprovincialis. Marine Ecology
Mieszkowska, N., Leaper, R., Moore, P., Kendall, M.A., Burrows, M.T., Lear, D., Poloczanska, E.S., Hiscock, K.,
Moschella, P.S., Thompson, R.C., Herbert, R.J., Laffoley, D., Baxter, J., Southward, A.J. & Hawkins, S.J.
2005. Assessing and Predicting the Influence of Climatic Change Using Intertidal Rocky Shore Biota.
Occasional Publications 20. Plymouth, U.K.: Marine Biological Association of the United Kingdom.
Milliman, J.D. 1991. Flux and fate of fluvial sediment and water in coastal seas. In Ocean Margin Processes
in Global Change, R.F.C. Mantoura (ed.). Chinchester, U.K.: John Wiley & Sons, 69–89.
Mitchell, B.G. & Holm-Hansen, O. 1991. Observations and modeling of the Antarctic phytoplankton crop in
relation to mixing depth. Deep-Sea Research Part A-Oceanographic Research Papers 38, 981–1007.
Modarressie, R., Rick, I.P. & Bakker, T.C.M. 2006. UV matters in shoaling decisions. Proceedings of the
Royal Society B 273, 849–854.
Moller, A.P., Flensted-Jensen, E. & Mardal, W. 2006. Rapidly advancing laying date in a seabird and the
changing advantage of early reproduction. Journal of Animal Ecology 75, 657–665.
Moore, L.R., Goericke, R. & Chisholm, S.W. 1995. Comparative physiology of Synechococcus and Prochlo-
rococcus: influence of light and temperature on growth, pigments, fluorescence and absorptive prop-
erties. Marine Ecology Progress Series 116, 259–275.
Moorthy, P. & Kathiresan, K. 1997. Influence of ultraviolet-B radiation on photosynthetic and biochemical
characteristics of a mangrove Rhizophora apiculata. Photosynthetica 34, 465–471.
Moorthy, P. & Kathiresan, K. 1998. Effects of UV-B irradiance on biomass and uptake of nutrients in mangrove
seedlings of Rhizophora apiculata (Rhizophorales: Rhizophoraceae). Indian Journal of Marine Sci-
ences 27, 239–242.
Mouritsen, K.N., Tompkins, D.M. & Poulin, R. 2005. Climate warming may cause a parasite-induced collapse
in coastal amphipod populations. Oecologia 146, 476–483.
Mrosovsky, N., Bass, A., Corliss, L.A., Richardson, J.I. & Richardson, T.H. 1992. Pivotal beach temperatures
for Hawksbill turtles nesting in Antigua. Canadian Journal of Zoology 70, 1920–1925.
Mumby, P.J. 2006. Connectivity of reef fish between mangroves and coral reefs: algorithms for the design of
marine reserves at seascape scales. Biological Conservation 128, 215–222.
Mumby, P.J., Edwards, A.J., Arias-Gonzalez, J.E., Lindeman, K.C., Blackwell, P.G., Gall, A., Gorczynska,
M.I., Harborne, A.R., Pescod, C.L., Renken, H., Wabnitz, C.C.C. & Llewellyn, G. 2004. Mangroves
enhance the biomass of coral reef fish communities in the Caribbean. Nature 427, 533–536.
Negri, A., Volhardt, C., Humphrey, C., Heyward, A., Jones, R., Eaglesham, G. & Fabricius, K. 2005. Effects
of the herbicide diuron on the early life history stages of coral. Marine Pollution Bulletin 51, 370–383.
469
Norkko, A., Thrush, S.F., Hewitt, J.E., Cummings, V.J., Norkko, J., Ellis, J.I., Funnell, G.A., Schultz, D. &
MacDonald, I. 2002. Smothering of estuarine sandflats by terrigenous clay: the role of wind-wave
disturbance and bioturbation in site-dependent macrofaunal recovery. Marine Ecology Progress Series
234, 23–41.
O’Brien, C.M., Fox, C.J., Planque, B. & Casey, J. 2000. Climate variability and North Sea cod. Nature 404, 142.
Ochieng, C.A. & Erftemeijer, P.L.A. 2002. Phenology, litterfall and nutrient resorption in Avicennia marina
(Forssk.) Vierh in Gazi Bay, Kenya. Trees Structure and Function 16, 167–171.
O’Hara, T.D. & Poore, G.C.B. 2000. Patterns of distribution for southern Australian marine echinoderms and
decapods. Journal of Biogeography 27, 1321–1335.
Okey, T.A., Vargo, G.A., Mackinson, S., Vasconcellos, M., Mahmoudi, B. & Meyer, C.A. 2004. Simulating
community effects of sea floor shading by plankton blooms over the West Florida Shelf. Ecological
Modelling 172, 339–359.
Oliver, J. & Babcock, R.C. 1992. Aspects of the fertilization ecology of broadcast spawning corals: sperm
dilution effects and in situ measurements of fertilization. Biological Bulletin 183, 409–417.
Orr, J.C., Fabry, V.J., Aumont, O., Bopp, L., Doney, S.C., Feely, R.A., Gnanadesikan, A., Gruber, N., Ishida,
A., Joos, F., Key, R.M., Lindsay, K., Maier-Reimer, E., Matear, R., Monfray, P., Mouchet, A., Najjar,
R.G., Plattner, G.K., Rodgers, K.B., Sabine, C.L., Sarmiento, J.L., Schlitzer, R., Slater, R.D., Totterdell,
I.J., Weirig, M.F., Yamanaka, Y. & Yool, A. 2005. Anthropogenic ocean acidification over the twenty-
first century and its impact on calcifying organisms. Nature 437, 681–686.
Palenik, B., Price, N.M. & Morel, F.M.M. 1991. Potential effects of UV-B on the chemical environment of
marine organisms: a review. Environmental Pollution 70, 117–130.
Pandolfi, J.M., Jackson, J.B., Baron, N., Bradbury, R.H., Guzman, H.M., Hughes, T.P., Kappel, C.V., Micheli,
F., Ogden, J.C., Possingham, H.P. & Sala, E. 2005. Are U.S. coral reefs on the slippery slope to slime?
Science 308, 1742–1743.
Parkinson, R.W., Delaune, R.D. & White, J.R. 1994. Holocene sea-level rise and the fate of mangrove forests
within the wider Caribbean region. Journal of Coastal Research 10, 1077–1086.
Parmesan, C., Ryrholm, N., Stefanescu, C., Hill, J.K., Thomas, C.D., Descimon, H., Huntley, B., Kaila, L.,
Kullberg, J., Tammaru, T., Tennent, W.J., Thomas, J.A. & Warren, M. 1999. Poleward shifts in
geographical ranges of butterfly species associated with regional warming. Nature 399, 579–583.
Parmesan, C. & Yohe, G. 2003. A globally coherent fingerprint of climate impacts across natural systems.
Nature 421, 37–42.
Pedersen, M.F. & Hansen, P.J. 2003a. Effects of high pH on a natural marine planktonic community. Marine
Pedersen, M.F. & Hansen, P.J. 2003b. Effects of high pH on the growth and survival of six marine heterotrophic
protists. Marine Ecology Progress Series 260, 33–41.
Pemberton, D. & Gales, R. 2004. Australian fur seals (Arctocephalus pusillus doriferus) breeding in Tasmania:
population size and status. Wildlife Research 31, 301–309.
Perez, T., Garrabou, J., Sartoretto, S., Harmelin, J.G., Francour, P. & Vacelet, J. 2000. Mass mortality of
marine invertebrates: an unprecedented event in the Northwestern Mediterranean. Comptes Rendus
de l’Academie des Sciences Serie III — Sciences de la Vie 323, 853–865.
Perry, A.L., Low, P.J., Ellis, J.R. & Reynolds, J.D. 2005. Climate change and distribution shifts in marine
fishes. Science 308, 1912–1915.
Peters, R.H. 1983. Size structure of the plankton community along the trophic gradient of Lake Memphremog.
Canadian Journal of Fisheries and Aquatic Sciences 40, 1770–1778.
Philippart, C.J.M., van Aken, H.M., Beukema, J.J., Bos, O.G., Cadee, G.C. & Dekker, R. 2003. Climate-
related changes in recruitment of the bivalve Macoma balthica. Limnology and Oceanography 48,
2171–2185.
Phillips, J.A. 2001. Marine macroalgal biodiversity hotspots: why is there high species richness and endemism
in southern Australian marine benthic flora? Biodiversity and Conservation 10, 1555–1577.
Pittock, A.B., Walsh, K. & McInnes, K. 1996. Tropical cyclones and coastal inundation under enhanced
greenhouse conditions. Water Air and Soil Pollution 92, 159–169.
470
Polacheck, T., Hobday, A., West, G., Bestley, S. & Gunn, J. 2006. Comparison of east-west movements of
archival tagged southern bluefin tuna in the 1990s and early 2000s. Prepared for the CCSBT 7th
Meeting of the Stock Assessment Group (SAG7) and the 11th Meeting of the Extended Scientific
Committee (ESC11) 4–11 September, and 12–15 September 2006, Tokyo, Japan. CCSBT-ESC/0609/28.
Polovina, J.J., Balazs, G.H., Howell, E.A., Parker, D.M., Seki, M.P. & Dutton, P.H. 2004. Forage and migration
habitat of loggerhead (Caretta caretta) and olive ridley (Lepidochelys olivacea) sea turtles in the
central north Pacific Ocean. Fisheries and Oceanography 13, 36–51.
Polovina, J.J., Haight, W.R., Moffitt, R.B. & Parrish, F.A. 1995. The role of benthic habitat, oceanography
and fishing on the population dynamics of the spiny lobster, Panulirus marginatus (Decapoda, Pali-
nuridae), in the Hawaiian archipelago. Crustaceana 68, 203–212.
Polovina, J.J., Mitchum, G.T., Graham, N.E., Craig, M.G., Demartini, E.E. & Flint, E.N. 1994. Physical and
biological consequences of a climate event in the central North Pacific. Fisheries Oceanography 3, 15–21.
Poore, G.C.B. 2001. Biogeography and diversity of Australia’s marine biota. In The State of the Marine
Environment Report for Australia Technical Annex: 1. The Marine Environment, L.P. Zann & P. Kailola
(eds). Department of the Environment, Sport and Territories, Ocean Rescue 2000 Program. Townsville,
Queensland, Australia: Great Barrier Reef Marine Park Authority, 75–84.
Porteiro C., Jacobson L.D., Rothschild B., De Oliveira J.A.A., Sanchez R.P., Barange M., Serra R., Cisneros-
Mata M.A., Uriarte A., Félix-Uraga R., Wada T., Hunter J.R., Kim J.Y., Matsuura Y. & Ñiquen M
2001. Surplus production, variability, and climate change in the great sardine and anchovy fisheries.
Canadian Journal of Fisheries and Aquatic Sciences 58,1891-1903.
Porter, J.W. & Meier, O.W. 1992. Quantification of loss and change in Floridian reef coral populations.
American Zoologist 32, 625–640.
Pörtner, H.O., Langenbuch, M. & Reipschläger, A. 2004. Biological impact of elevated ocean CO2 concen-
trations: lessons from animal physiology and earth history. Journal of Oceanography 60, 705–718.
Preen, A.R., Long, W.J.L. & Coles, R.G. 1995. Flood and cyclone related loss, and partial recovery, of more
than 1000 km2 of seagrass in Hervey Bay, Queensland, Australia. Aquatic Botany 52, 1–2.
Prowse, T.A.A. & Pile, A.J. 2005. Phenotypic homogeneity of two intertidal snails across a wave exposure
gradient in South Australia. Marine Biology Research 1, 176–185.
Przeslawski, R., Davis, A.R. & Benkendorff, K. 2004. Effects of ultraviolet radiation and visible light on the
development of encapsulated molluscan embryos. Marine Ecology Progress Series 268, 151–160.
Przeslawski, R., Davis, A.R. & Benkendorff, K. 2005. Synergistic effects associated with climate change and
the development of rocky shore molluscs. Global Change Biology 11, 515–522.
Ralph, P.J. 1998. Photosynthetic response of laboratory cultured Halophila ovalis to thermal stress. Marine
Raven, J., Caldeira, K., Elderfield, H., Hoegh-Guldberg, O., Liss, P, Riebesell, U., Shepherd, J., Turley, C. &
Watson, A. 2005. Ocean Acidification due to Increasing Atmospheric Carbon Dioxide. London: Royal
Society Special Report.
Reed, D.C. & Foster, M.S. 1984. The effects of canopy shading on algal recruitment and growth in a giant
kelp forest. Ecology 65, 937–948.
Rhoads, D.C., Boesch, D.F., Zhican, T., Fengshan, X., Liqiang, H. & Nilsen, K.J. 1985. Macrobenthos and
sedimentary facies on the Changjiang delta platform and adjacent continental shelf, East China Sea.
Continental Shelf Research 4, 189–213.
Rich, H.W. & Morel, F.M.M. 1990. Availability of well-defined iron colloids to the marine diatom Thalassiosira
weissflogii. Limnology and Oceanography 35, 652–662.
Richardson, A.J. & Schoeman, D.S. 2004. Climate impact on plankton ecosystems in the Northeast Atlantic.
Science 305, 1609–1612.
Richardson, A.J. & Verheye, H.M. 1998. The relative importance of food and temperature to copepod egg
production and somatic growth in the southern Benguela upwelling system. Journal of Plankton
Research 20, 2379–2399
Richer de Forges, B., Koslow, J.A. & Poore, G.C.B. 2000. Diversity and endemism of the benthic seamount
fauna in the southwest Pacific. Nature 405, 944–947.
471
Ridgway, K.R. & Condie, S.A. 2004. The 5500 km-long boundary flow off western and southern Australia.
Journal of Geophysical Research 109, C04017.
Ridgway, K.R. & Dunn, J.R. 2003. Mesoscale structure of the mean East Australian Current System and its
relationship with topography. Progress in Oceanography 56, 189–222.
Ridgway, K.R. & Godfrey, J.S. 1997. Seasonal cycle of the East Australian Current. Journal of Geophysical
Research 102, 22,921–22,936.
Riebesell, U., Zondervan, I., Rost, B., Tortell, P.D., Zeebe, R.E. & Morel, F.M.M. 2000. Reduced calcification
of marine plankton in response to increased atmospheric CO2. Nature 407, 364–367.
Rijstenbil, J.W., Coelho, S.M. & Eijsackers, M. 2000. A method for the assessment of light-induced oxidative
stress in embryos of fucoid algae via confocal laserscan microscopy. Marine Biology 137, 763–774.
Rivadeneira, M.M. & Fernandez, M. 2005. Shifts in southern endpoints of distribution in rocky intertidal
species along the south-eastern Pacific coast. Journal of Biogeography 32, 203–209.
Rivalan, P., Prévot-Julliard, A.C., Choquet, R., Pradel, R., Jacquemin, B. & Girondot, M. 2005. Trade-off
between current reproductive effort and delay to next reproduction in the leatherback sea turtle.
Oecologia 145, 564–574.
Roberts, J.M., Long, D., Wilson, J.B., Mortensen, P.B. & Gage, J.D. 2003. The cold-water coral Lophelia
pertusa (Scleractinia) and enigmatic seabed mounds along the north-east Atlantic margin: are they
related? Marine Pollution Bulletin 46, 7–20.
Roberts, J.M., Wheeler, A.J. & Freiwald, A. 2006. Reefs of the deep: the biology and geology of cold-water
coral ecosystems. Science 312, 543–547.
Robinson, R.A., Learmonth, J.A., Hutson, A.M., Macleod, C.D., Sparks, T.H., Leech, D.I., Pierce, G.J.,
Rehfisch, M.M. & Crick, H.Q.P. 2005. Climate change and migratory species. BTO Research Report
414, British Trust for Ornithology, Norfolk, U.K., 85–88.
Roemmich, D. & McGowan, J. 1995. Climatic warming and the decline of zooplankton in the California
Current. Science 267, 1324–1326.
Roessig, J.M., Woodle, C.M., Cech, J.J., Jr. & Hansen, L.J. 2004. Effects of global climate change on marine
and estuarine fishes and fisheries. Reviews in Fish Biology and Fisheries 14, 251–275.
Rogers, C.S. 1990. Responses of coral reefs and reef organisms to sedimentation. Marine Ecology Progress
Series 62, 185–202.
Rogers, K., Wilton, K.M. & Saintilan, N. 2006. Vegetation change and surface elevation dynamics in estuarine
wetlands of southeast Australia. Estuarine Coastal and Shelf Science 66, 559–569.
Root, T.L., Price, J.T., Hall, K.R., Schneider, S.H., Rosenzweigk, C. & Pounds, J.A. 2003. Fingerprints of
global warming on wild animals and plants. Nature 421, 57–60.
Rose, G.A. 2005a. On distributional responses of North Atlantic fish to climate change. ICES Journal of
Rose, G.A. 2005b. Capelin (Mallotus villosus) distribution and climate: a sea ‘canary’ for marine ecosystem
change. ICES Journal of Marine Science 62, 1524–1530.
Ross, G.J.B., Burbidge, A.A., Brothers, N., Canty, P., Dann, P., Fuller, P.J., Kerry, K.R., Norman, F.I.,
Menkhorst, P.W., Pemberton, D., Shaughnessy, G., Shaughnessy, P.D., Smith, G.C., Stokes, T. &
Tranter, J. 2001. The status of Australia’s seabirds. In The State of the Marine Environment Report
for Australia Technical Annex: 1 The Marine Environment, L.P. Zann & P. Kailola (eds). Department
of the Environment, Sport and Territories, Ocean Rescue 2000 Program. Townsville, Queensland,
Australia: Great Barrier Reef Marine Park Authority, 167–182.
Rothlisberg, P., Staples, D., Poiner, I. & Wolanski, E. 1998. The possible impact of the greenhouse effect on
commercial prawn populations in the Gulf of Carpentaria. In Greenhouse: Planning for Climate
Change, G.I. Pearman (ed.). Melbourne, Victoria, Australia: CSIRO Publications, 216–227.
Russell, F.S. & Colman, J.S. 1935. The zooplankton, 4. The occurrence and seasonal distribution of the
Tunicata, Mollusca and Coelenterata (Siphonophora). Scientific Reports of the Great Barrier Reef
Expedition 2, 203–276.
Sagarin, R.D., Barry, J.P., Gilman, S.E. & Baxter, C.H. 1999. Climate-related change in an intertidal community
over short and long time scales. Ecological Monographs 69, 465–490.
Saintilan, N. & Williams, R.J. 1999. Mangrove transgression into saltmarsh environments in south-east
Australia. Global Ecology and Biogeography 8, 117–124.
472
Sammarco, P.W. & Andrews, J.C. 1988. Localised dispersal and recruitment in Great Barrier Reef Corals: the
Helix experiment. Science 239, 1422–1424.
Sarmiento, J.L., Slater, R., Barber, R., Bopp, L., Doney, S.C., Hirst, A.C., Kleypas, J., Matear, R., Mikolajewicz,
U., Monfray, P., Soldatov, V., Spall, S.A. & Stouffer, R. 2004. Response of ocean ecosystems to
climate warming. Global Biogeochemical Cycles 18, GB3003.
Sato, K., Matsuzawa, Y., Tanaka, H., Bando, T., Minamikawa, S., Sakamoto, W. & Naito, Y. 1998. Internesting
intervals for loggerhead turtles, Caretta caretta, and green turtles, Chelonia mydas, are affected by
temperature. Canadian Journal of Zoology 76, 1651–1662.
Scheibling, R.E. & Hennigar, A.W. 1997. Recurrent outbreaks of disease in sea urchins Strongylocentrotus
droebachiensis in Nova Scotia: evidence for a link with large-scale meteorologic and oceanographic
events. Marine Ecology Progress Series 152, 155–165.
Schiel, D.R., Steinbeck, J.R. & Foster, M.S. 2004. Ten years of induced ocean warming causes comprehensive
changes in marine benthic communities. Ecology 85, 1833–1839.
Schiermeier, Q. 2005. Cleaner skies leave global warming forecasts uncertain. Nature 435, 135.
Schippers, P., Lürling, M. & Scheffer, M. 2004. Increase of atmospheric CO2 promotes phytoplankton
productivity. Ecology Letters 7, 446–451.
Schwartz, M.D., Ahas, R. & Aasa, A. 2006. Onset of spring starting earlier across the Northern Hemisphere.
Global Change Biology 12, 343–351.
Sebens, K.P. 2002. Energetic constraints and size gradients in intertidal and subtidal marine invertebrates.
Integrative and Comparative Biology 42, 853–861.
Seddon, S., Connolly, R.M. & Edyvane, K.S. 2000. Large-scale seagrass dieback in northern Spencer Gulf,
South Australia. Aquatic Botany 66, 297–310.
Sedwick, P.N., DiTulliio, G.R., Hutchins, D.A., Boyd, P.W., Griffiths, F.B., Crossley, A.C., Trull, T.W. &
Quéguiner, B. 1999. Limitation of algal production by iron and silicic acid deficiency in the Australian
subantarctic region. Geophysical Research Letters 26, 2865–2868.
Serrão, E.A., Pearson, G., Kautsky, L. & Brawley, S.H. 1996. Successful external fertilization in turbulent
environments. Proceedings of the National Academy of Sciences of the United States of America 93,
5286–5290.
Setlow, R.B. & Setlow, J.K. 1962. Evidence that ultraviolet-induced thymine dimers in DNA cause biological
damage. Proceedings of the National Academy of Sciences of the United States of America 48,
1250–1257.
Shick, J.M., Lesser, M.P. & Jokiel, P.L. 1996. Effects of ultraviolet radiation on corals and other coral reef
organisms. Global Change Biology 2, 527–545.
Short, F.T. & Neckles, H.A. 1999. The effects of global climate change on seagrasses. Aquatic Botany 63,
169–196.
Siebeck, U.E. 2004. Communication in coral reef fish: the role of ultraviolet colour patterns in damselfish
territorial behaviour. Animal Behaviour 68, 273–282.
Simkanin, C., Power, A., Myers, A., McGrath, D., Southward, A.J., Mieszkowska, N., Leaper, R. & O’Riordan,
R. 2005. Using historical data to detect temporal changes in the abundances of intertidal species on
Irish shores. Journal of the Marine Biological Association of the United Kingdom 85, 1329–1340.
Simpson, C.J. 1991. Mass spawning of corals on Western Australian reefs and comparisons with the Great
Barrier Reef. Journal of the Royal Society of Western Australia 74, 85–92.
Sims, D.W., Genner, M.J., Southward, A.J. & Hawkins, S.J. 2001. Timing of squid migration reflects North
Atlantic climate variability. Proceedings of the Royal Society of London B 268, 2607–2611.
Sims, D.W., Wearmouth, V.J., Genner, M.J., Southward, A.J. & Hawkins, S.J. 2004. Low-temperature-driven
early spawning migration of a temperate marine fish. Journal of Animal Ecology 73, 333–341.
Smart, J. & Gill, J.A. 2003. Climate change and the potential impact on breeding waders in the U.K. Wader
Study Group Bulletin 100, 80–85.
Smayda, T.J. 1976. Phytoplankton processes in mid-Atlantic nearshore and shelf waters and energy-related
activities. In Effects of Energy-Related Activities on the Atlantic Continental Shelf, B Manowitz (ed.).
Report No. 50484. Brookhaven National Laboratory, Upton, New York, U.S., 70–95.
Smith, J.R., Fong, P. & Ambrose, R.F. 2006. Dramatic declines in mussel bed community diversity: response
to climate change? Ecology 87, 1153–1161.
473
Smith, R. & Baker, K. 1979. Penetration of UV-B and biologically effective dose-rates in natural waters.
Journal of Photochemistry and Photobiology 50, 459–468.
Snelgrove, P.V.R. & Smith, C.R. 2002. A riot of species in an environmental calm: the paradox of the species-
rich deep sea. Oceanography and Marine Biology: An Annual Review 40, 311–342.
Solow, A.R., Bjorndal, K.A. & Bolten, A.B. 2002. Annual variation in nesting numbers of marine turtles: the
effect of sea surface temperature on re-migration intervals. Ecology Letters 5, 742–746.
Southward, A.J., Hawkins, S.J. & Burrows, M.T. 1995. Seventy years’ observations of changes in distribution
and abundance of zooplankton and intertidal organisms in the western English Channel in relation to
rising sea temperature. Journal of Thermal Biology 20, 127–155.
Spalding, H., Foster, M.S. & Heine, J.N. 2003. Composition, distribution, and abundance of deep-water
(>30 m) macroalgae in central California. Journal of Phycology 39, 273–284.
Staples, D.J. 1980. Ecology of juvenile and adolescent banana prawns, Penaeus merguiensis, in a mangrove
estuary and adjacent off-shore area of the Gulf of Carpentaria. 1. Immigration and settlement of
postlarvae. Australian Journal of Marine and Freshwater Research 31, 635–652.
Staples, D.J., Dall, W. & Vance, D.J. 1982. Banana prawn catch prediction. Cleveland, Australia: CSIRO
Marine Laboratory Research Report 156, 31–41.
Staples, D.J. & Vance, D.J. 1986. Emigration of juvenile banana prawns Penaeus merguiensis from a mangrove
estuary and recruitment to offshore areas in the wet-dry tropics of the Gulf of Carpentaria, Australia.
Steneck, R.S., Graham, M.H., Bourque, B.J., Corbett, D., Erlandson, J.M., Estes, J.A. & Tegner, M.J. 2002. Kelp
forest ecosystems: biodiversity, stability, resilience and future. Environmental Conservation 29, 436–459.
Stewart, H.L. & Carpenter, R.C. 2003. The effects of morphology and water flow on photosynthesis of marine
macroalgae. Ecology 84, 2999–3012.
Stoner, D.S. 1992. Vertical distribution of a colonial ascidian on a coral reef: the roles of larval dispersal and
life-history variation. American Naturalist 139, 802–824.
Surman, C.A. & Wooller, R.D. 2000. Seabirds off the south-western coast of Australia. Emu 100, 312–317.
Swanson, A.K. & Druehl, L.D. 2000. Differential meiospore size and tolerance of ultraviolet light stress within
and among kelp species along a depth gradient. Marine Biology 136, 657–664.
Tegner, M.J., Dayton, P.K., Edwards, P.B. & Riser, K.L. 1996. Is there evidence for long-term climatic change
in southern California kelp forests? California Cooperative Oceanic Fisheries Investigations Reports
37, 111–126.
Thessen, A.E., Dortch, Q., Parsons, M.L. & Morrison, W. 2005. Effect of salinity on Pseudo-nitzschia species
(Bacillariophyceae) growth and distribution. Journal of Phycology 41, 21–29.
Thomas, C.D. & Lennon, J.J. 1999. Birds extend their ranges northwards. Nature, 399, 213.
Thomas, L.P., Moore, D.R. & Work, R.C. 1961. Effects of hurricane Donna on the turtle grass beds of Biscayne
Bay, Florida. Bulletin of Marine Science 11, 191–197.
Thresher, R.E. 2002. Solar correlates of Southern Hemisphere mid-latitude climate variability. International
Journal of Climatology 22, 901–915.
Thresher, R.E., Andrews, N. & Rowling, K. 1996. Interim Monitoring of the 1995 Eastern Gemfish Spawning
Run 95/039. Hobart, Tasmania: CSIRO Division of Fisheries/NSW Fisheries.
Thresher, R.E., Harris, G.P., Gunn, J.S. & Clementson, L.A. 1989. Planktonic production pulses and episodic
settlement of a temperate marine fish. Nature 341, 641–642.
Thresher, R.E., Nichols, P.D., Gunn, J.S., Bruce, B.D. & Furlani, D.M. 1992. Evidence for microbial production
based on seagrass detritus supporting a planktonic food chain. Limnology and Oceanography 37,
1754–1758.
Thresher, R., Rintoul, S.R., Koslow, J.A., Weidman, C., Adkins, J. & Proctor, C. 2004. Oceanic evidence of
climate change in southern Australia over the last three centuries. Geophysical Research Letters 31,
Art. No. L07212.
Thrush, S.F., Hewitt, J.E., Cummings, V., Ellis, J.I., Hatton, C., Lohrer, A. & Norkko, A. 2004. Muddy waters:
elevating sediment input to coastal and estuarine habitats. Frontiers in Ecology and the Environment
2, 299–306.
Thrush, S.F., Hewitt, J.E., Herman, P.M.J. & Ysebaert, T. 2005. Multi-scale analysis of species-environment
relationships. Marine Ecology Progress Series 302, 13–26.
474
Thrush, S.F., Hewitt, J.E., Norkko, A., Cummings, V.J. & Funnell, G.A. 2003a. Macrobenthic recovery
processes following catastrophic sedimentation on estuarine sandflats. Ecological Applications 13,
1433–1455.
Thrush, S.F., Hewitt, J.E., Norkko, A., Nicholls, P.E., Funnell, G.A. & Ellis, J.I. 2003b. Habitat change in
estuaries: predicting broad-scale responses of intertidal macrofauna to sediment mud content. Marine
Tortell, P.D., DiTullio, G.R., Sigman, D.M. & Morel, F.M.M. 2002. CO2 effects on taxonomic composition
and nutrient utilization in an equatorial Pacific phytoplankton assemblage. Marine Ecology Progress
Series 236, 37–43.
Travis, J.M.J. 2002. Climate change and habitat destruction: a deadly anthropogenic cocktail. Proceedings of
the Royal Society B 270, 467–473.
Tsuchiya, M. 1983. Mass mortality in a population of the mussel Mytilus edulis L. caused by high temperature
on rocky shores. Journal of Experimental Marine Biology and Ecology 66, 101–111.
Twilley, R.R. & Chen, R. 1998. A water budget and hydrology model of a basin mangrove forest in Rookery
Bay, Florida. Marine and Freshwater Research 49, 309–323.
Tyagi, A.P. 2004. Precipitation effect on flowering and propagule setting in mangroves of the family Rhizo-
phoraceae. Australian Journal of Botany 52, 789–798.
Vadas, R.L. & Steneck, R.S. 1988. Zonation of deep-water benthic algae in the Gulf of Maine. Journal of
Valentine, J.P. & Johnson, C.R. 2004. Establishment of the introduced kelp Undaria pinnatifida following
dieback of the native macroalga Phyllospora comosa in Tasmania, Australia. Marine and Freshwater
Research 55, 223–230.
Vance, D., Staples, D. & Kerr, J. 1985. Factors affecting year-to-year variation in the catch of banana prawns
(Penaeus merguiensis) in the Gulf of Carpentaria, Australia. Journal du Conseil International pour
l’Exploration de la Me 42, 83–97.
Vance, D.J., Haywood, M.D.E., Heales, D.S., Kenyon, R.A. & Loneragan, N.R. 1998. Seasonal and annual
variation in abundance of postlarval and juvenile banana prawns Penaeus merguiensis and environ-
mental variation in two estuaries in tropical northeastern Australia: a 6-year study. Marine Ecology
Venrick, E.L., McGowan, J.A., Cayan, D.R. & Hayward, T.L. 1987. Climate and chlorophyll a: long-term
trends in the Central North Pacific Ocean. Science 238, 70–72.
Vilchis, L.I., Tegner, M.J., Moore, J.D., Friedman, J.D., Friedman, C.S., Riser, K.L., Robbins, T.T. & Dayton,
P.K. 2005 Ocean warming effects on growth reproduction and survivorship of southern California
abalone. Ecological Applications 15, 469–480.
Votier, S.C., Hatchwell, B.J., Beckerman, A., McCleery, R.H., Hunter, F.M., Pellatt, J., Trinder, M. & Birkhead,
T.R. 2005. Oil pollution and climate have wide-scale impacts on seabird demographics. Ecology
Letters 8, 1157–1164.
Wahl, M., Molis, M., Davis, A., Dobretsov, S., Durr, S.T., Johansson, J., Kinley, J., Kirugara, D., Langer, M.,
Lotze, H.K., Thiel, M., Thomason, J.C., Worm, B. & Ben-Yosef, D.Z. 2004. UV effects that come
and go: a global comparison of marine benthic community level impacts. Global Change Biology 10,
1962–1972.
Walker, D., Dennison, W. & Edgar, G. 1999. Status of Australian seagrass research and knowledge. In Seagrass
in Australia: Strategic Review and Development of an R & D Plan, A.J. Butler & P. Jernakoff (eds).
Collingwood, Australia: CSIRO Publishing, 1–24.
Walker, D.I., Kendrick, G.A. & McComb, A.J. 1988. The distribution of seagrass species in Shark Bay, Western
Australia, with notes on their ecology. Aquatic Botany 30, 305–317.
Walker, D.I & Prince, R.I.T. 1987. Distribution and biogeography of seagrass species on the northwest coast
of Australia. Aquatic Botany 29, 19–32.
Walker, T.A. 1992. A record crested tern Sterna bergii colony and concentrated breeding by seabirds in the
Gulf of Carpentaria. Emu 92, 152–156.
Walsh, K. & Pittock, A.B. 1998. Potential changes in tropical storms, hurricanes and extreme rainfall events
as a result of climate change. Climatic Change 39, 199–213.
475
Walsh, K.J.E., Betts, H., Church, J., Pittock, A.B., McInnes, K.L., Jackett, D.R. & McDougall, T.J. 2004.
Using sea-level rise projections for urban planning in Australia. Journal of Coastal Research 20,
586–598.
Walther, G.R., Berger, S. & Sykes, M.T. 2005. An ecological ‘footprint’ of climate change. Proceedings of
the Royal Society Series B 272, 1427–1432.
Walther, G.R., Post, E., Convey, E., Menzel, A., Parmesan, C., Beebee, T.J.C., Fromentin, J.M., Hoegh-
Guldberg, O. & Bairlein, F. 2002. Ecological responses to recent climate change. Nature 416, 389–395.
Wängberg, S.A., Garde, K., Gustavson, K. & Selmer, J.S. 1999. Effects of UV-B radiation on marine
phytoplankton communities. Journal of Plankton Research 21, 147–166
Ward, J.R. & Lafferty, K.D. 2004. The elusive baseline of marine disease: are diseases in ocean ecosystems
increasing? PLOS Biology 2, 0542–0547.
Ward, T.M., Hoedt, F., McLeay, L., Dimmlich, W.F., Kinloch, M., Jackson, G., McGarvey, R., Rogers, P.J. &
Jones, K. 2001. Effects of the 1995 and 1998 mass mortality events on the spawning biomass of
sardine, Sardinops sagax, in South Australian waters. ICES Journal of Marine Science 58, 865–875.
Ware, D.M. 1995. A century and a half of change in the climate of the North East Pacific. Fisheries
Webster, P.J., Holland, G.J., Curry, J.A. & Chan, H.R. 2005. Changes in tropical cyclone number, duration
and intensity in a warming environment. Science 309, 1844–1846.
Weishampel, J.F., Bagley, D.A. & Ehrhart, L.M. 2004. Earlier nesting by loggerhead sea turtles following sea
surface warming. Global Change Biology 10, 1424–1427.
Wellington, G.M. & Fitt, W.K. 2003. Influence of UV radiation on the survival of larvae from broadcast-
spawning reef corals. Marine Biology 143, 1185–1192.
Wells, F.W. 1985 Zoogeographical importance of tropical marine mollusc species at Rottnest Island, W.A.
Western Australian Naturalist 16, 40–45.
Welsford, D.C. & Lyle, J.M. 2003. Redbait (Emmelichthys nitidus): a synopsis of fishery and biological data.
TAFI Technical Report 20, Tasmania, Australia.
West, R.J. & Larkum, A.W.D. 1979. Leaf productivity of the seagrass, Posidonia australis, in eastern Australian
waters. Aquatic Botany 7, 57–65.
White, W.B., Gloersen, K.A., Marsac, F. & Tourre, Y.M. 2004. Influence of coupled Rossby waves on primary
productivity and tuna abundance in the Indian Ocean. Journal of Oceanography 60, 531–541.
Whitehead, P.J. & Saalfeld, K. 2000. Nesting phenology of magpie geese (Anseranas semipalmata) in
monsoonal northern Australia: responses to antecedent rainfall. Journal of Zoology 251, 495–508.
Wiencke, C., Roleda, M.Y., Gruber, A., Clayton, M.N. & Bischof, K. 2006. Susceptibility of zoospores to
UV radiation determines upper depth distribution limit of Arctic kelps: evidence through field exper-
iments. Journal of Ecology 94, 455–463.
Wilhelm, C., Bida, J., Domin, A., Hilse, C., Kaiser, B., Kesselmeier, J., Lohr, M. & Müller, A.M. 1997.
Interaction between global climate change and the physiological responses of algae. Photosynthetica
33, 491–503.
Wilkinson, C.R. 1999. Global and local threats to coral reef functioning and existence: review and predictions.
Marine and Freshwater Research 50, 867–878.
Wilkinson, C. (ed.) 2004. Status of Coral Reefs of the World: 2004. Townsville, Queensland, Australia:
Australian Institute of Marine Science, 301 pp.
Williams, A., Gowlett-Holmes, K. & Althaus, F. 2006. Biodiversity Survey of the Seamounts and Slopes of
the Norfolk Ridge and Lord Howe Rise (NORFANZ). Final Report to the National Oceans Office,
April 2006. Hobart, Australia: CSIRO Marine and Atmospheric Research.
Willis, J.K., Roemmich, D. & Cornuelle, B. 2003. Combining altimetric height with broadscale profile data
to estimate steric height, heat storage, subsurface temperature, and sea-surface temperature variability.
Journal of Geophysical Research — Oceans 108, Art. No. 3292.
Witherington, B.E. 2002. Ecology of neonate loggerheads inhabiting lines of downwelling near a Gulf Stream
front. Marine Biology 140, 843–853.
Wolanski, E. & Chappell, J. 1996. The response of tropical Australian estuaries to a sea level rise. Journal of
Marine Systems 7, 267–279.
476
Wolfe, D.W., Schwartz, M.D., Lakso, A.N., Otsuki, Y., Pool, R.M. & Shaulis, N.J. 2005. Climate change and
shifts in spring phenology of three horticultural woody perennials in northeastern U.S.A. International
Journal of Biometeorology 49, 303–309.
Womersley, H.B.S. & Edmonds, S.J. 1958, A general account of the intertidal ecology of south Australian
coasts. Australian Journal of Marine and Freshwater Research 9, 217-260.
Woodroffe, C. 1992. Mangrove sediments and geomorphology. In Tropical Mangrove Ecosystems, Coastal
and Estuarine Studies No. 41, A.I. Robertson & D.M. Alongi (eds), Coastal and Estuarine Studies
41, 7–42.
Worm, B., Lotze, H.K. & Myers, R.A. 2003. Predator diversity hotspots in the blue ocean. Proceedings of
the National Academy of Sciences of the United States of America 100, 9884–9888.
Worrest, R.C., van Dyke, H. & Thomson, B.E. 1978. Impact of enhanced simulated solar ultraviolet radiation
upon a marine community. Journal of Photochemistry and Photobiology B 27, 471–478.
Yntema, C.L. & Mrosovsky, N. 1982. Critical periods and pivotal temperatures for sexual differentiation in
loggerhead sea turtles. Canadian Journal of Zoology 60, 1012–1016.
Zacherl, D., Gaines, S.D. & Lonhart, S.I. 2003. The limits to biogeographical distributions: insights from the
northward range extension of the marine snail, Kelletia kelletii (Forbes, 1852). Journal of Biogeog-
raphy 30, 913–924.
Zagaglia, C.R. & Stech, J.L. 2004. Remote sensing data and longline catches of yellowfin tuna (Thunnus
albacares) in the equatorial Atlantic. Remote Sensing of the Environment 93, 267–281.
Zagarese, H.E. & Williamson, C.E. 2000. Impact of UV radiation on zooplankton and fish. In The Effects of
UV Radiation in the Marine Environment. S.J. de Mora et al. (eds) Cambridge, UK: Cambridge
University Press, 279-309.
Zagarese, H.E. & Williamson, C.E. 2001. The implications of solar UV radiation exposure for fish and fisheries.
Fish and Fisheries 2, 250–260.
Zamzow, J.P. 2004. Effects of diet, ultraviolet exposure, and gender on the ultraviolet absorbance of fish mucus
and ocular structures. Marine Biology 144, 1057–1064.
Zann, L.P. 2000. The eastern Australian region: a dynamic tropical/temperate biotone. Marine Pollution
Bulletin 41, 188–203.
Zein-Eldin, Z.P. & Renaud, M.L. 1986. Inshore environmental effects on brown shrimp Penaeus aztecus and
white shrimp P. setiferus populations in coastal waters, particularly of Texas. Marine Fisheries Review
48, 9–19.
Zimmerman, R.C., Korhs, D.G., Steller, D.L. & Alberte, R.S. 1997. Impacts of CO2 enrichment on productivity
and light requirements of eelgrass. Plant Physiology 115, 599–607.
Zimmerman, R.C. & Robertson, D.L. 1985. Effects of El Niño on hydrography and growth of the giant kelp,
Macrocystis pyrifera, at Santa Catalina Island, California. Limnology and Oceanography 30,
1298–1302.
Zou, D.H. & Gao, K.S. 2002. Photosynthetic responses to inorganic carbon in Ulva lactuca under aquatic
and aerial states. Acta Botanica Sinica 44, 1291–1296.
Zou, D.H. & Gao, K.S. 2005. Ecophysiological characteristics of four intertidal marine macroalgae during
emersion along Shantou coast of China, with a special reference to the relationship of photosynthesis
and CO2. Acta Oceanologica Sinica 24, 105–113.
477
APPENDIX: THE CSIRO MK3 CLIMATE SYSTEM MODEL

The CSIRO Mk3 Climate System Model (CSIRO Mk3) is a state-of-the-art climate model that
represents all the major components of the Earth’s climate system: atmosphere, land surface, sea
ice and oceans. A detailed description of the CSIRO Mk3 is given in Gordon et al. (2002) and
summarised below. The CSIRO Mk3 is ranked with the top international models.
CSIRO Mk3 model simulations are included in the Intergovernmental Panel on Climate Change
(IPCC) Fourth Assessment Report (IPCC 2007). The Fourth Assessment Report assesses scientific,
technical and socioeconomic information relevant for the understanding of climate change, its
potential impacts, and the options for adaptation and mitigation. An essential component of this
report is climate change projections of the impact of various scenarios of future levels of greenhouse
gases on the earth’s climate system. The CSIRO Mk3 simulations are an important contributor to
these climate projections. For climate projection present in this study we use the IPCC SRES A2
greenhouse gas emission scenario, which projects atmospheric CO2 levels of 536 ppm by 2050.
The CSIRO Mk3 atmospheric module has a spectral T63 horizontal grid (~1.875° latitude by
1.875° longitude) with 18 vertical levels (hybrid sigma-pressure vertical co-ordinate). The atmo-
spheric model includes a comprehensive cloud microphysical parameterisation and convection
parameterisation, which are linked via the detrainment of liquid and frozen water at the cloud top.
Atmospheric moisture advection (vapour, liquid and frozen) is carried out by the semi-Lagrangian
method. This module includes the direct radiative forcing of sulphate on atmospheric albedo.
The CSIRO Mk3 land surface scheme uses six layers of moisture and temperature with a
vegetation canopy. The scheme uses multiple soil (9) and vegetation (12) types and includes a
three-layer snow model.
The sea-ice module incorporates a dynamical-thermodynamic polar ice model that includes a
variable fraction of leads. The CSIRO Mk3 ocean model is based upon the Modular Ocean Model
version 2.2 (MOM2.2) of the Geophysical Fluid Dynamics Laboratory (GFDL) model. The oceanic
component has horizontal resolution of ~0.9375° latitude by 1.875° longitude. For every atmo-
spheric grid point there are two ocean points in the meridional direction, which allows for the
atmospheric model and ocean model subcomponents to have matching land-sea masks. There are
31 levels in the vertical, with the spacing of the levels gradually increasing with depth, from 10 m
at the surface to 400 m at depth. The ocean model includes a parameterisation of mixing of tracers
based on the formulation of Griffies et al. (1998) and improved vertical mixing in the tropical Pacific.
For the climate change projection in this manuscript we use the Mk 3.5 version of the climate
system model which has a greatly improved simulation of the Southern Ocean from the original
Mk 3 climate system model.
478
β (θ)
s γ β (θ) = β (θ)
β (θ) [m−1sr−1] β (θ) [m−1sr−1]; t = 2
200
A B C 100
105 104 50 102
50
20
20 102 10
102 101
10
5
5 100
2
2 10−2 100
1
Particle size D (µm)

10−2
1
0.5
50931_C009.fm Page 549 Tuesday, May 1, 2007 6:09 PM
Phase shift parameter ρ
0.5
m = 1.05 + i0.01 m = 1.05 + i0.01 10−4 m = 1.05 + i0.01 0.2 10−1
10−5
0.2
200
D E F 300
105 200
50 103 100 102
20 50
102
10 20 101
101
5 10
2 5 100
10−2 10−1
Particle size D (µm)

Phase shift parameter ρ
1 2
0.5 1 10−1
m = 1.17 + i0.0001 m = 1.17 + i0.0001 10−3 m = 1.17 + i0.0001
0.2 10−5 0.5
0 30 60 90 120 150 180 0 30 60 90 120 150 180 0 30 60 90 120 150 180
Scattering angle θ [deg] Scattering angle θ [deg] Scattering angle θ [deg]
Colour Figure 4 (Clavano, Boss & Karp-Boss) The volume scattering function for spheres, β (θ) (A, D), and for equal-volume spheroids, β (θ) with aspect ratio s/t = 2
(B, E). The ratio between the two (i.e., the bias denoted as γβ(θ) is presented in panels C and F. The primary y-axis for each plot represents variation in particle size, D[µm],
while the secondary y-axis represents variation in the phase shift parameter, ρ (scale found on C and F). Results are for two different types of particles: phytoplankton-like
particles with m = 1.05 + i0.01 (A, B, C) and inorganic-like particles with m = 1.17 + i0.0001 (D, E, F). Values for spheroids have been obtained using the T-matrix method for
D ≤ 10 µm and by the ray tracing method for D ≥ 40 µm. No solution is available for 10 < D < 40 µm (white regions in B, C, E and F).
Day of July 1995, West Sound, Orcas Island, Washington

26 27 28 29

20
10
0
midnight noon midnight noon midnight noon
−75 −70 −65 −60 −55 −50 −45 −40 −35 −30 −25
Volume backscattering strength at 265 kHz (dB)
Day of August 2002, Damariscotta River, Maine

19 20 21 22
10
0
midnight noon midnight noon midnight
0 200 400 600 800 1000 1200 1400 1600

Biovolume concentration (mm3 m−3)
Colour Figure 5 (Jumars) For legend see Jumars Figure 5 (p. 120).
26°S
73° 72° 71°
28°S 28° 28°
30°S
29° 29°
73°W 72°W 71°W 30°

30°
73° 72° 71°

60 0 60 120 180 Kilometers
Colour Figure 3 (Thiel) For legend see Thiel Figure 3 (p. 207).
20 20 20
15 15 15
−25
10 10 10
5 5 5
−30
20 22 24 2 20 22 24 2 20 22 24 2
26–27 July 27–28 July 28–29 July
−35
20 20 20
15 15 15 −40
10 10 10
Volume backscattering strength (dB)

5 5 5 −45
20 22 24 2 20 22 24 2 20 22 24 2
29–30 July 30–31 July 30 July–1 August −50
20 20 20
15 15 15 −55
10 10 10
−60
5 5 5
2 4 6 8 2 4 6 8 2 4 6 8
27 July 28 July 29 July −65
20 20 20
−70
15 15 15
10 10 10
−75
5 5 5
2 4 6 8 2 4 6 8 2 4 6 8
30 July 31 July 1 August
Hour of day
Colour Figure 6 (Jumars) For legend see Jumars Figure 6 (p. 120).
10°
Exmouth Torres Strait
Gulf Darwin Gulf of
Indian Scott
Carpentaria
Ocean Reef Cape
York Great Barrier Reef
20°
Hervey Bay Pacific Ocean
Australia
Shark Bay
Brisbane Moreton Bay
Houtman Abrolhos
30° Islands
Hawkesbury Estuary
Perth Adelaide Sydney Botany Bay
Albany
Melbourne
Corner Inlet New Zealand
Bass Strait Tasman Sea
40° Tasmania
Hobart
Southern Ocean Tasmanian Seamounts

Marine Reserve
50°
110° 120° 130° 140° 150° 160° 170° 180° 190°
Colour Figure 1 (Poloczanska et al.) Map of Australia indicating the locations discussed in the text. The
200 nm EEZ for Australia is marked by the dashed line, and the 200 m depth contour by the solid line.
110°00’E 120°00’E 130°00’E 140°00’E 150°00’E

10°00’S N 10°00’S
1b
Darwin
1c?
2a
1a Kimberlays 2c
Cairns
Burketown
20°00’S Port Hedland Northern 20°00’S
Territory Mackey 3
Queensland
Australia
Western Australia
Brisbane
30°00’S South Australia 30°00’S
New South
5a Perth Ceduna Wales
Esperance Sydney
2b Adelaide 2d
Canberra
Victoria ACT
Melbourne 5b
4 2d
40°00’S 40°00’S
Tasmania 2d
6 Hobart
110°00’E 120°00’E 130°00’E 140°00’E 150°00’E
Colour Figure 3 (Poloczanska et al.) Phytoplankton provinces around Australia. In northern shelf waters
westwards from Torres Strait tropical diatom species dominate, with slight regional differences in relative
abundances and absolute biomass (1a-c). The shallow waters of the Great Barrier Reef region (3) are dominated
by fast-growing nano-sized diatoms. The deeper waters of the Indian Ocean and the Coral Sea are characterised
by a tropical oceanic flora (2a and 2c, respectively) that is dominated by dinoflagellates and follows the
Leeuwin Current (2b) and the East Australia Current and its eddies (2d). South-eastern coastal waters harbour
a temperate phytoplankton flora (4) with seasonal succession of different diatom and dinoflagellate commu-
nities. Waters south of the tropical and temperate phytoplankton provinces are characterised by an oceanic
transition flora (5a,b) that communicates to the subantarctic phytoplankton province (6) and is highly variable
in extent. The phytoplankton provinces are associated with surface water masses and the zooplankton fauna
likely shows a similar pattern (Figure prepared by G.M. Hallegraeff for CSIRO and National Oceans Office).
10°N 35 10°N 10 10°N 260

240
30 8
0° 0° 0° 220
6 200
25
10°S 10°S 10°S
180
4
20 160
20°S 20°S 2 20°S
140
15
0 120
30°S 30°S 30°S
10 100
−2
80
40°S 40°S 40°S
5 60
−4
50°S 50°S 50°S 40
0 −6
20
60°S −5 60°S −8 60°S 0
20.0 cm/s
10°N 2.6 10°N 6 10°N 10

2.4 5 0
0° 2.2 0° 4 0° −10
2 3 −20
10°S
1.8
10°S
2
10°S −30
−40
1.6 1
20°S 20°S 20°S −50
1.4 0
−60
1.2 −1
30°S 30°S 30°S −70
1 −2
−80
40°S
0.8 40°S
−3 40°S −90
0.6 −4 −100
50°S 0.4 50°S −5 50°S −110
0.2 −6 −120
60°S 0 60°S −7 60°S −130
5.00 cm/s
Colour Figure 5 (Poloczanska et al.) Simulated annual means of SST (°C) with annual mean surface currents
(cm/s) (left), annual mean zonal winds (m/s) (middle), and mixed layer depth (m) (right). In the middle panels,
westerly wind direction is denoted by positive sign, easterly wind direction by negative sign. Top row: 1990s,
bottom row: difference between 1990s and 2070s.
10ºN 8.16 10ºN 7

8.14 6.5
0º 0º
8.12
6
10ºS 8.1 10ºS
5.5
8.08
20ºS 20ºS 5
8.06
30ºS 8.04 30ºS 4.5

8.02
4
40ºS 8 40ºS
3.5
7.98
50ºS 50ºS
7.96 3
60ºS 7.94 60ºS 2.5

60
80
10
12
14
16
18
60
80
10
12
14
16
18
ºE
ºE
0º
0º
0º
0º
0º
ºE
ºE
0º
0º
0º
0º
0º
E
E
10ºN −0.09 10ºN −0.5
−0.1 −0.6
0º 0º
−0.11 −0.7
10ºS −0.12 10ºS −0.8
−0.13 −0.9
20ºS 20ºS
−0.14 −1
30ºS −0.15 30ºS −1.1
40ºS −0.16 40ºS −1.2

−0.17 −1.3
50ºS 50ºS
−0.18 −1.4
60ºS −0.19 60ºS −1.5

60
80
10
12
14
16
18
60
80
10
12
14
16
18
ºE
ºE
0º
0º
0º
0º
0º
ºE
ºE
0º
0º
0º
0º
0º
E
Colour Figure 6 (Poloczanska et al.) Simulated annual means of pH (left) and aragonite saturation state
(right). Top row: 1990s, bottom row: difference between 1990s and 2070s.
10ºN 280 10ºN 15 10ºN 260

260 14 240
0º 0º 0º 220
240 12
10 200
10ºS 220 10ºS 10ºS
180
8
200 20ºS 160
20ºS 6 20ºS
180 140
30ºS 4
30ºS 30ºS 120
160
2
100
40ºS 140 40ºS 0 40ºS
80
120 −2
50ºS 60
50ºS 50ºS
100 −4 40
60ºS 80 60ºS −6 60ºS 20
60
80
10
12
14
16
18
60
80
10
12
14
16
18
60
80
10
12
14
16
18
ºE
ºE
0º
0º
0º
0º
0º
ºE
ºE
0º
0º
0º
0º
0º
ºE
ºE
0º
0º
0º
0º
0º
E
E
E
E
E
E
E
E
E
E
E
E
10ºN 100 10ºN 3 10ºN 28
2.5 26
80
0º 0º 0º 24
2
60 22
10ºS 10ºS 1.5 10ºS 20
40
1 18
20ºS 20 20ºS 20ºS
0.5 16
0 0 14
30ºS 30ºS 30ºS
−0.5 12
−20 10
40ºS 40ºS −1 40ºS
−40 8
−1.5
50ºS 50ºS 50ºS 6
−60 −2 4
60ºS −80 60ºS −2.5 60ºS 2
60
80
10
12
14
16
18
60
80
10
12
14
16
18
60
80
10
12
14
16
18
ºE
ºE
0º
0º
0º
0º
0º
ºE
ºE
0º
0º
0º
0º
0º
ºE
ºE
0º
0º
0º
0º
0º
E
E
E
E
E
E
E
E
E
E
E
E
Colour Figure 7 (Poloczanska et al.) Simulated annual means of downward solar radiation at the ocean
surface (W/m2) (left), precipitation minus evaporation (mm/d) (middle), and sea-level height anomaly due to
upper ocean stratification relative to 2000 m (cm) (right). Top row: 1990s, bottom row: difference between
1990s and 2070s.
Present AL
TROPIC
RATE
TEMPE
Future AL
TROPIC
RATE
TEMPE
beach island turtle – female tropical fish tropical coral

tropical
kelp forest turtle – male temperate fish bleached coral
mangroves
tropical calcifying
seagrass dugong jellyfish carbonate rock plankton
Colour Figure 9 (Poloczanska et al.) Hypothetical Southern Hemisphere marine coastline and coastal
waters ranging from low latitudes in the north to high latitudes in the south under present climate (top) and
in the future under global warming scenario (bottom). As temperatures rise, species’ distributions shift further
south. The range of tropical and subtropical species extends to temperate latitudes while temperate species in
the south decline. Rising temperatures and ocean acidification stress coral reefs leading to frequent coral
bleaching and an increase in mortality while rapid sea level rise inundates the coral reefs. Ocean acidification
also leads to the decline of calcifying plankton such as pteropods and coccolithophores. Rising sea-level
encroaches on the mainland and on offshore islands. The sex ratio of marine turtle hatchlings, which is
determined by ambient nest temperatures, is skewed in the future as warming produces more females.
50931_Idx1.fm Page 479 Thursday, May 10, 2007 8:34 PM
AUTHOR INDEX
References to complete articles are given in bold type, references to bibliographic lists are in regular type.
A Aguilar, R. See Guerra, C., 318

See Luna-Jorquera, G., 325
Aadnesen, A. See Kaartvedt, S., 131 Aguilar-Rosas, L.E., 74
Aasa, A. See Menzel, A., 469 Aguilar-Rosas, R. See Aguilar-Rosas, L.E., 74
See Schwartz, M.D., 473 Aguilera, C. See Martínez, G., 326
Aaser, H., 124 Aguirre, G. See Sielfeld, W., 337
Abal, E.G., 453 Ahas, R. See Menzel, A., 469
See Carruthers, T.J.B., 457 See Schwartz, M.D., 473
Abarca, A. See Uriarte, I., 340 Ahrens, M. See Lohrer, A.M., 467
See von Brand, E., 343 Ahumada, R., 305
Abbiati, M. See Airoldi, L., 394 See Bernal, P.A., 307
See Moschella, P.S., 402 See González, H.E., 317
Abbott, I.A., 74 Aiken, C., 305
Abello, H.U., 124 Ainsworth, E.A., 453
See Taylor, L.H., 137 Airamè, S., 393
Aberg, P. See Jonsson, P.R., 465 Airoldi, L., 345–405, 393, 394
See Airoldi, L., 394 See Bacchiocchi, F., 394
See Moschella, P.S., 402 See Benedetti-Cecchi, L., 395
Able, K.W. See Beck, M.W., 394 See Bulleri, F., 396
See Nemerson, D.M., 133 See Moschella, P.S., 402
Abraham, E.R. See Boyd, P.W., 456 Akaboshi, S. See Illanes, J.E., 321
Abreu-Grobois, F.A. See Bowen, B.W., 455 Akano, T. See Gao, K., 461
Akin, T.B. See Mattison, J.E., 83
Aburto, J., 304
Aksnes, D.L. See Kaartvedt, S., 131
See Stotz, W., 338
Alarcón, E., 305
Aburto-Oropeza, O. See Sala, E., 334
See Wolff, M., 344
Aceves, H. See Langdon, C., 466
Alarcón, G. See Montecino, V., 327
Acha, E.M. See Schiariti, A., 135
Alarcón, R. See Acuña, E., 304
Acuña, E., 304, 305
Albaina, A. See Bonnet, D., 455
See Arancibia, H., 305
Alberte, R.S. See Zimmerman, R.C., 477
See Castilla, J.C., 310
Albertsson, J., 124
See Sielfeld, W., 336
Albrecht, A. See Schories, D., 403
See Thiel, M., 199–344
Alcalde, O. See Neill, P.E., 329
Acuña, E.H. See Villarroel, J.C., 343
Alcock, A., 190
Acunto, S. See Piazzi, L., 403 Alder, J. See Agardy, T., 393
Adam, P., 393 Alderete, W. See Majluf, P., 325
Adami, M.L., 74 Alexander, S. See Roughgarden, J., 333
Adams, D.C. See Rosenberg, M.S., 171 Alheit, J., 305
Ades, P.K. See Keatley, M.R., 465 Aliaga, B. See Bernal, P.A., 307
Adey, W.H., 453 Allan, R., 190
Adkins, J. See Thresher, R., 474 Allee, W.C., 166
Adnan, N.A. See Loneragan, N.R., 467 Allen, D.M., 125
Adnyana, W., 453 See Johnson, W.S., 130
Aedo, D. See Santelices, B., 85 Allen, H.D., 394
See Santelices, B., 334, 335 Allen, J.A., 166
Aerts, J. See Kabat, P.W., 400 See Rex, M.A., 170
Aerts, K. See Engel, A., 460 Allen, J.H. See Connor, D.W., 396
Afanasyev, V. See Phillips, R.A., 332 Allen, J.R.L., 394
Agardy, T., 393 Allison, G., Russell, R., 171
Agegian, C.R. See Cowen, R.K., 76 Alm, M.B. See Hutchins, D.A., 321
Aguayo, A. See Sielfeld, W., 336 Alm-Kübler, K. See Menzel, A., 469
Aguilar, A., 305 Aloia, A. See Colombini, I., 311
Aguilar, M., 305 Alongi, D.M., 394, 453
See González, S.A., 318 Alonso Vega, J.M. See Thiel, M., 199–344
479
AUTHOR INDEX
Alpers, W., 190 Apostolaki, E.T. See Karakassis, I., 169

Althaus, F. See Williams, A., 476 Arana, P., 305
Altizer, S. See Harvell, C.D., 463 See Palma, S., 331
Alvarado, J.L., 305 Arancibia, A. See Acuña, E., 304
See Castilla, J.C., 310 Arancibia, H., 305
Alvarez, E. See Diaz-Almela, E., 458 See Medina, M., 326
Alveal, K., 305 See Moloney, C.L., 327
Amado, N. See Sielfeld, W., 336 See Neira, S., 330
Amakasu, K., 125 Araneda, A. See Escribano, R., 314
Amann, R. See Kuypers, M.M., 323 Araneda, S.E. See Vargas, C.A., 341
Amarasekare, P. See Leibold, M.A., 323 Arata, J., 305
See Micheli, F., 327 See Báez, P., 306
Ambarsari, I. See Brown, B.E., 191 See Moreno, C.A., 328
Ambrose, R.F. See Smith, J.R., 473 Araya, B. See Simeone, A., 337
Amoureux, L. See Cornet, M., 127 Araya, M. See Medina, M., 326
Amsler, C.D., 74 Archaux, F., 454
See Reed, D.C., 85, 333 Archer, D. See Kleypas, J.A., 466
An, Z.S. See Jickells, T.D., 465 Arcos, D. See Cáceres, M., 309
Anandrai, A. See Perissinotto, R., 134 See Letelier, J., 324
Andelman, S. See Leslie, H., 324, 400 Arcos, D.F. 305
See Lubchenco J., 169 See Cubillos, L.A., 312
See Meir, E., 327 See Peterson, W.T., 331
See Possingham, H., 403 See Vargas, C.A., 341
Andelman, S.J., 166 Ardiwijaya, R.L. See Baird, A.H., 190
Andersen, K.K. See Jickells, T.D., 465 Argua, Y. See Gao, K., 461
Andersen, R.A. See Coyer, J.A., 76 Ariani, A.P., 125
Anderson, A.N., 166 See Wittman, K.J., 138
Anderson, B.S., 74 Arias, A.M. See Drake, P., 128
Anderson, D.M. See Moy, C.M., 329 Arias-Gonzalez, J.E. See Mumby, P.J., 469
Anderson, E.K., 74 Arlhac, D. See Salen-Picard, C., 334
Anderson, J.M. See Moschella, P.S., 402 Armstrong, R.A. See Detres, Y., 458
Anderson, R.J. See Stegenga, H., 86 Arnaud, J. See Brunet, M., 126
Anderson, T.W., 74 Arnold, K.E., 74
See Reed, D.C., 85 Arntz, W., 305, 306
Andersson, A. See Jonzén, N., 465 See Brante, A., 308
Andrade, M.J. See Acuña, E., 305 See Fernández, M., 316
See Villarroel, J.C., 343 See Palma, M., 331
Andrade, Y.N. See Camus, P.A., 309 See Rosenberg, R., 333
Andrew, N.L., 453 See Thiel, M., 199–344
See Schiel, D.R., 86 Arntz, W.E., 306
Andrews, J.C. See Sammarco, P.W., 473 See Peña T.S., 331
Andrews, N. See Thresher, R.E., 474 See Tarazona, J., 339
Andrews, N.L., 74 Aro, E. See Flinkman, J., 128
Angel, A., 305 Aron, A. See Villegas, A., 343
Anggoro A.W. See Baird, A.H., 190 Aronson, R.B., 125
Anghera, M. See Reed, D.C., 85 Arriaza, B.T. See Santoro, C.M., 335
Anraku, M. See Clutter, R.I., 126 Arsenault, D.J., 454
Ansaloni, I. See Simonini, R., 404 Artemyev, A.V. See Both, C., 455
Antezana, T., 125, 305 Aruga, Y. See Gao, K.S., 461
See Hückstädt, L.A., 320 Asada, K. See Gao, K., 461
Anthony, E.J., 394 Asano, E., 33
Anthony, K.R.N., 190 Asencio, G. See Duarte, W.E., 313
Antia, A.N., 454 See Moreno, C.A., 328
Antoine, D. See Morel, A., 35 Asensi, A., 74
Antolic, B. See Meinesz, A., 401 Ashjian, C.J. See Lawson, G.L., 132
Antonov, J.I. See Conkright, M.E., 311 Asmus, H. See Asmus, R.M., 306
Apel, M., 125 Asmus, R.M., 306
480
AUTHOR INDEX
Asprey, K.W. See Syvitski, J.P.M., 36 Badalamenti, F., 394

Astier, J.M. See Meinesz, A., 401 See Milazzo, M., 402
Astill, K. See Booth, D.T., 455 Baden, S., 394
Astoreca, R. See Montecino, V., 327 See Pihl, L., 403
Astorga, A., 306 Baden, S.P. See Boström, C., 395
Astorga, M. See Galleguillos, R., 316 Badola, R., 454
Astthorsson, O.S., 125 Báez, P., 306
Atkinson, L. See Yannicelli, B., 344 Baeza, J.A., 306
Atkinson, L.P., 306 See Fernández, M., 316
Atkinson, M.J. See Langdon, C., 466 Bagley, D.A. See Weishampel, J.F., 476
Atkinson, R.J.A. See Hall-Spencer, J.M., 399 Bahamonde, N. See Castilla, J.C., 310
Attramadai, Y.G., 125 Bahamondes, I. See Castilla, J.C., 76
Attrill, M.J., 166 Bainbridge, R., 125
See Foggo, A., 167 Baird, A.H., 190
Attwood, C.G., 74 See Hughes, T.P., 464
Au, W.W.L. See Benoit-Bird, K.J., 125 Bairlein, F. See Walther, G.R., 476
Audzijonyte, A., 125 Bais, A. See McKenzie, R.L., 468
Augenstein, E.W., 74 Bak, R.P.M., 190
Aumont, O. See Bopp, L., 455 Baker, A.C. See Drohan, A.F., 459
See Orr, J.C., 470 Baker, A.R. See Jickells, T.D., 465
Aure, J. See Erga, S.R., 460 Baker, K. See Smith, R., 474
Aursland, K. See Erga, S.R., 460 Bakker, D.C.E. See Boyd, P.W., 456
Austen, M.C., 166 Bakker, J.P. See Wolters, M., 405
See Widdicombe, S., 172 Bakker, T.C.M. See Modarressie, R., 469
Australia Bureau of Statistics, 454 Bakun, A., 306
Australian Greenhouse Office, 454 See Cury, P., 312
Autio, H. See Kangas, P., 400 See Weeks, S.J., 343
Auyang, S.Y., 125 Balanda, M.J. See Thiel, M., 339
Avaria S., 306
Balazs, G.H. See Bowen, B.W., 455
Avellanal, M.H. See Jaramillo, E., 321, 322
See Polovina, J.J., 471
Avendaño, M., 306
Balbontín, F., 306
See Moragat, D., 328
Baldó, F. See Drake, P., 128
Avila, M., 75, 306
Baldwin, R. See Kaufmann, R.S., 131
See Correa, J.A., 312
Balech, E., 307
See Hoffmann, A.J., 81, 320
Balestri, E. See Diaz-Almela, E., 458
Aviléz, O., 306
Ball, B.J., 394
Avishai, N. See Baruch, R., 454
Ball, I. See Leslie, H., 400
Axelsson, L. See Lundälv, T., 401
See Possingham, H., 403
Aydin, K., 33
Ball, I.R. See Leslie, H., 324
Ayukai, T. See McKinnon, A.D., 468
Ballesteros, E., 394
Azeiteiro, U.M.M., 125
See Boudouresque, C.F., 395
Azuma, N. See Takahashi, K., 136
See Delgado, O., 396
See Thibaut, T., 404
Balmford, A., 307, 394
B Baltazar, M. See Gallardo, V.A., 316
Baardseth, E., 75 See Gutiérrez, D., 319
Baba, J., 33 See Roa, R., 333
Babcock, E.A. See Majluf, P., 325 Bamber, R.N., 125
Babcock, R., 454 Bancroft, W.J., 454
Babcock, R.C. See Cole, R.G., 457 Bando, T. See Sato, K., 473
See Poloczanska, E.S., 407–478, 454 Bannon, S.M. See MacLeod, C.D., 468
See Shears, N.T., 336 Barange M. See Porteiro C., 471
See Oliver, J., 470 Barber, P.W. See Geller, P.E., 34
Bacchiocchi, F., 394 Barber, R. See Sarmiento, J.L., 473
See Moschella, P.S., 402 Barber, R.T., 125
Bacescu, M., 125 See Codispoti, L.A., 311
Bach, C.S., 454 Barbera, C., 394
481
AUTHOR INDEX
Barbieri, E.S., 454 Beauvais, L., 190

Barbieri, M. See Yáñez, E., 344 Beck, M.W., 307, 394, 395
Barbieri, M.A. See Bertrand, A., 307 See Airoldi, L., 345–405, 394
See Silva, C., 337 Becker, J., 307
See Yáñez, E., 344 Beckerman, A. See Votier, S.C., 475
Barilotti, D.C., 75 Beckley, L.E., 75
Barker, D. See Procter, T.D., 35 Bedo, A.W. See Longhurst, A.R., 132
Barker, N.P. See Teske, P.R., 339 Beebee, T.J.C. See Walther, G.R., 476
Barmawidjaja, D., 394 Beer, S. See Beardall, J., 454
Barnabe, G., 394 Behrenfeld, M., 455
Barnard, A.H. See Twardowski, M.S., 36 Behrens, M.D., 75
Barnes, D.J. See Lough, J.M., 192 Beker, J.B. See Connor, D.W., 396
See Lough, J.M., 467 Bell, A.M. See Duke, N.C., 459
See McNeil, B.I., 469 Bell, S.S. See Fonseca, M.S., 460
Barnes, J.H., 454 Bellantoni, D. See Peterson, W.T., 331
Barnett, H. See Langdon, C., 466 Bellwood, D.R. See Hughes, T.P., 464
Baron, J. See Moragat, D., 328 Belmonte, G. See Fanelli, G., 398
Baron, N. See Pandolfi, J.M., 470 Belsher, T., 75
Barr, N. See Wing, B.L., 138 See Meinesz, A., 401
Barrales, H.L., 75 Bence, J.R. See Nisbet, R.M., 84
Barrera, J.C. See Sala, E., 334 Benedetti-Cecchi, L., 395
Barreto, R. See Godfrey, M.H., 461 Benkendorff, K. See Przeslawski, R., 471
Barrett, N.S., 397 Ben Maiz, N. See See Boudouresque, C.F., 395
See Edgar, G.J., 459 Bennett, A.F. See MacNally, R., 169
Barria, I. See Lagos, N., 323 Bennett, W.A. See Kimmerer, W.J., 131
Barry, J.P., 454 Benninger, L.K. See Muñoz, P., 329
See Sagarin, R.D., 472 Benoit-Bird, K.J., 125
Barry, T.M. See Lesser, M.P., 467 Benthien, A. See Engel, A., 460
Barth, J.A. See Grantham, B.A., 318 Ben-Yosef, D.Z. See Wahl, M., 475
Bartlett, K.P. See Kunze, E., 131 Berasategui, A.D. See Schiariti, A., 135
Barton, E.D. See Hill, A.E., 320 Bergametti, G. See Jickells, T.D., 465
Bartsch, I., 394 Berge, J.A., 455
Baruch, R., 454 Berger, S. See Walther, G.R., 476
Bascompte, J. See Micheli, F., 327 Berger, U. See Grimm, V., 129
Bass, A. See Mrosovsky, N., 469 Berger, W.H. See Jackson, J.B.C., 399
Basterretxea, G. See Marbà, N., 401 Berges, J. A. See Wilson, S., 405
Bastias, H. See Thiel, M., 199–344 Berghahn, R., 395
Bastreri, D. See Calliari, D., 126 Berghe, E.V. See Costello, M.J., 167
Bathmann, U. See Antia, A.N., 454 Bergman, A. See Zhivotovsky, L.A., 138
Battisti, D.S. See Zhang, Y., 344 Bergmann, T. See Boss, E., 33
Baumgartner, T. See Montecino, V., 327 Bergstrom, L. See Eriksson, B.K., 398
Baumgartner, T.R. See Halpin, P.M., 319 Berkelmans, R. See Done, T.J., 459
Baums, I.B., 307 Berkenbusch, K. See Lohrer, A.M., 467
Bavestrello, G., 394 Berlow, E.L. See Menge, B.A., 327
See Cicogna, F., 396 Bernal, M., 307
Baxter, C.H. See Barry, J.P., 454 See Simeone, A., 337
See Sagarin, R.D., 472 Bernal, P. See González, H.E., 317
Baxter, J. See Mieszkowska, N., 469 See Iriarte, J.L., 321
Beamish, R.J., 454 See Llanos, A., 324
See Burkett, V., 456 See Llanos-Rivera, A., 324
Beardall, J., 454 Bernal, P.A., 307
Beardsley, G.F. See Carder, K.L., 33 See Castro, L.R., 310
Beare, D.J., 454 See González, H.E., 318
Beaudouin, J., 125 Bernard, S. See Quirantes, A., 36
Beaufort, L. See Engel, A., 460 Bernardes, C. See Cunha, M.R., 127
Beaugrand, G., 454 Bernhard, J.M. See Gooday, A.J., 318
See Bonnet, D., 455 Bernhard, M. See Jeftic, L., 400
See Kirby, R.R., 466 Bernstein, B.L., 75
Beaumont, L.J., 454 Berríos, V. See Sielfeld, W., 337
482
AUTHOR INDEX
Bertasi, F. See Martin, D., 401 See Morales, C.E., 328

Bertignac, M. See Lehodey, P., 467 See Thomas, A.C., 339
Bertness, M.D. See Wolters, M., 405 See Uribe, E., 340
Bertrán, C. See Jaramillo, E., 322 See Valle-Levinson, A., 341
Bertrand, A., 307 Blanco, M. See Edding, M., 313
Bertrand, N. See Winkler, G., 138 Blanz, T. See Antia, A.N., 454
Bertrand, Y., 166 Blasco, F., 455
Bestley, S. See Polacheck, T., 471 Bleakley, C. See Kelleher, G., 400
Betts, H. See Walsh, K.J.E., 476 Bloomfield, A.L., 455
Betzer, P.R. See Carder, K.L., 33 Boaden, P.J.S. See Birkett, D.A., 395
Beucher, M. See Asensi, A., 74 Bobadilla, M., 307
Beukema, J.J. See Philippart, C.J.M., 470 See Santelices, B., 85
Beveridge, I. See Kunze, E., 131 Bodini, A. See Acuña, E., 305
Beyst, B., 125 Boehlert, G.W. See Koslow, J.A., 466
Bhargava, R.M.S. See Desai, B.N., 191 Boer, G.J. See Cubasch, U., 458
Bhattacharya, D. See Yoon, H.S., 88 Gregory, J.M., 462
Bi, D. See Cai, W., 456 Boero, F. See Fanelli, G., 398
Bianchi, C.N., 395 See Guidetti, P., 398
See Sandulli, R., 403 See Marcus, N.H., 325
Bickmore, B.R., 33 Boersma, D., 307
Bida, J. See Wilhelm, C., 476 Boesch, D.F. See Rhoads, D.C., 471
Bilham, R., 191 Boghen, A.D. See St-Hilaire, A., 136
Billot, C. See Faugeron, S., 315 Bogucki, D. See Stramski, D., 36
Binckley, C.A., 455 Bohm, G. See Yáñez, E., 344
Bird, E.C.F., 395 Bohren, C.F., 33
Bird, J., 307 Boisset, F. See Boudouresque, C.F., 395
Birkett, D.A., 395 Bokn, T.L., 395
Birkhead, T.R. See Votier, S.C., 475 Boland, J.W. See McArthur, L.C., 468
Birklund, J. See van Dalfsen, J. A., 405 Boland, W. See Maier, I., 83
Birowo, A.T. See Soegiarto, A., 194 Bollens, S.M., 455
Bischof, K., 307, 455 See Brown, H., 126
See Wiencke, C., 476 See Dean, A.F., 127
Bishop, J.D.D. See Gage, J., Boltaña, S. See Hinojosa, I., 320
Bissolli, P. See Menzel, A., 469 See Macaya, E.C., 83, 325
Bjerkeng, B. See Berge, J.A., 455 Bolten, A.B. See Solow, A.R., 474
Bjørgesæter, A. See Gray, J.S., 168 Bolton, J.J., 455
Bjorn, L.O. See Madronich, S., 325 See Stegenga, H., 86
See McKenzie, R.L., 468 Boltovskoy, E., 307
Bjorndal, K.A. See Jackson, J.B.C., 399 Bolvaran, A. See Sielfeld, W., 336
See Pandolfi, J.M., 402 Bombace, G., 395
See Solow, A.R., 474 Bomkamp, R.E., 307
Bjornsen, P.K. See Nielsen, T.G., 192 Bonaventura, R., 455
Blaauw, B. See Both, C., 455 Bondesan, M., 395
Black, K.P. See McShane, P.E., 469 Boni, L. See Carlsson, P., 456
Blackall, L.L. See Jones, R.J., 465 Bonnet, D., 455
Blackburn, S., 455 Bonsdorff, E. See Karlson, K., 400
Blackburn, S.I., 455 Bonvicini Pagliai, A.M. See Simonini, R., 404
Blackwell, B., 455 Boo, S.M. See Yoon, H.S., 88
Blackwell, P.G. See Mumby, P.J., 469 Boonruang, P. See Nielsen, T.G., 192
Blain, S.P. See Coale, K.H., 311 Boonyanate, P. See Chansang, H., 191
Blair, D., See Adnyana, W., 453 Boorman, L.A., 395
Blake, C. See Wilson, S., 405 Booth, D.T., 455
Blanchette, C.A., 75 Bopp, L., 455
See Menge, B.A., 327 See Orr, J.C., 470
Blanco, J.L., 307 See Sarmiento, J.L., 473
See Atkinson, L.P., 306 Borcard, D. See Legendre, P., 169
See Carr, M.E., 309 Bordehore, C. See Barbera, C., 394
See Castro, L.R., 311 Borg, J.A. See Barbera, C., 394
See Escribano, R., 315 Bortman, M.L. See Beck, M.W., 395
483
AUTHOR INDEX
Bortolus, A. See Orensanz, J.M., 134 Brand, L.E., 456

Bos, O.G. See Philippart, C.J.M., 470 Brandhorst, W., 307
Boschi, E.E. See Astorga, A., 306 Brandon M.A. See Brierley A.S., 126
Boss, E., 33 Brandt, A., 126
See Clavano, W.R., 1–38 Branstrator, D.K., 126
See Roesler, C.S., 36 Brante, A., 308
See Stramski, D., 36 See Fernández, M., 315
See Twardowski, M.S., 36 Braslavska, O. See Menzel, A., 469
Boström, C., 395 Brattegard, T., 126
See Worm, B., 405 See Fosså, J.H., 129
Both, C., 455 Brattstöm, H., 308
Botsford, L.W., 307 Braun, M. See Escribano, R., 314
See Jackson, J.B.C., 399 See Morales, C.E., 328
See Lockwood, D.R., 324 Bravo, L. See Fuenzalida, R., 316
See Wing, S.R., 88, 343 See Ramos, M., 333
Böttjer, D., 307 Brawley, S.H., 456
Botto-Mahan, C. See Ebensperger, L.A., 78, 313 See Serrão, E.A., 473
Boucher, G. See Lambshead, P.J.D, 169 Bray, M.J., 395
Bouchet, J.-M. See Cornet, M., 127 Bray, R.N., 75
Bouchet, P. See Costello, M.J., 167 Brearley, A. See Jernakoff, P., 465
See Rex, M.A., 170 Breck, E. See Leslie, H., 324
Boudouresque, C.F., 395 Breeman, A.M. See Peters, A.F., 331
See Meinesz, A., 401 Breen, P.A. See Druehl, L.D., 77
Boudreau, B.P., 126 Bregman, A. See Kelfkens, G., 465 465
Boulding, E.G., 455 Breslin, V.T. See Song, K.-H., 136
Boulter, S.L., 455 Brey, L. See Bischof, K., 307
Bourdeau, P. See Stanners, D., 404 Brey, T., 308
Bourge, I. See Gattuso, J.P., 461 See Arntz W.E., 306
Bourget, E. See Le Fèvre, J., 323 Bricaud, A., 33
Bourgois, T. See Remerie, T., 134 See Morel, A., 35
Bourque, B.J. See Jackson, J.B.C., 399 See Stramski, D., 36
See Lotze, H.K., 401, Briede, A. See Menzel, A., 469
See Steneck, R.S., 86, 404, 474 Brierley A.S., 126
Bowden, K.F., 307 Brinch, C. See Jonzén, N., 465
Bowen, B.W., 455 Brink, K.H. See Hill, A.E., 320
Bowen, J.L. See Valiela, I., 405 Brink, L. See Shanks, A.L., 86
Bowen, P. See Abal, E.G., 453 Briones, L., 308
Bowie, A.R. See Boyd, P.W., 456 Britton-Simmons, K. See Klinger, T., 322
Bowman, D.M.J.S., 456 Broccoli, A.J. See Rosenthal, Y., 85
Bowman, T.E. See Williams, A.B., 138 Brochier, T. See Guizien, K., 318
Bowyer, J. See Jones, R.J., 465 Broderick, A.C., 456
Boyd, P.W., 456 See Hays, G.C., 463
See Jickells, T.D., 465 Brodie, J., 456
See Sedwick, P.N., 473 See Devlin, M.J., 458
Boyer, T.P. See Conkright, M.E., 311 Broitman, B. See Navarrete, S.A., 329
Bozec, Y.M. See Moloney, C.L., 327 Broitman, B.R., 75, 308
Bozinovic, F. See Pulgar, J.M., 332 See Blanchette, C.A., 75
Bozzano, A., 126 See Halpern, B.S., 80
Bracher, D. See Farrell, T.M., 315 See Navarrete, S.A., 329
Bracken, M.E.S. See Menge, B.A., 327 Brokordt, K. See Martínez, G., 326
Bradbury, R.H. See Jackson, J.B.C., 399 See Thiel, M., 199–344
See Lotze, H.K., 401 Brokordt, K.B., 308
See Pandolfi, J.M., 402, 470 Brooks, N. See Jickells, T.D., 465
Brainard, R.E. See Benoit-Bird, K.J., 125 Brooks, T.M., 395
Braje, T.J. See Erlandson, J.M., 78 Brostoff, W.N., 75
Branch, G. See Boersma, D., 307 Brothers, N. See Bunce, A., 456
See Roberts, C.M., 333 See Ross, G.J.B., 472
Branch, G.M. See Beckley, L.E., 75 Browman, H.I., 395
See Bustamante, R.H., 75, 308 See Kouwenberg, J.H.M., 466
484
AUTHOR INDEX
Brown, A.C., 395 See Mieszkowska, N., 469

Brown, B.E., 173–194, 191 See Southward, A.J., 474
See Clarke, K.R., 191 Burt, M.D.B. See Jackson, C.J., 130
See Dunne, R.P., 191 Burton, H. See Lake, S., 132
See Satapoomin, U., 193 Burton, R.S., 308
See Scoffin, T.P., 193 Busch, M. See Lotze, H.K., 401
Brown, G.W. See MacNally, R., 169 Buschmann, A.H., 75, 308
Brown, H., 126 See Gutierrez, A., 80
Brown, J.H., 308 See Castilla, J.C., 310
Brown, M.T., 75 See Chopin, T., 311
See Chin, N.K.M., 76 See Graham, M.H., 39–88, 318
See Correa, J.A., 312 See Gutierrez, A., 319
See Nyman, M.A., 84 See Macaya, E.C., 83, 325
Brubaker, J. See Shanks, A.L., 86 See Munoz, V., 83, 329
Bruce, B.D. See Thresher, R.E., See Vásquez, J.A., 87, 341, 342
Bruhn, J., 456 See Vega, J.M.A., 88, 342
Bruland, K.W. See Hutchins, D.A., 321 Bustamante, R.H., 75, 308
Brumbaugh, D.R. See Harborne, A.R., 463 See Boersma, D., 307
Brunel, P., 126 See Roberts, C.M., 333
Brunet, M., 126 Bustos, C. See Landaeta, M.F., 323
Brunsting, A. See Latimer, P., 35 Butler, A. See Poloczanska, E.S., 407–478
Brzezinski, M.A. See Reed, D.C., 85 Butler, A.J. See Davis, A.R., 458
Bucarey, D.A. See Cubillos, L.A., 312 See Keough, M.J., 466
Buchanan, J.B., 126 Butman, C.A., 308
See Frid, C.L., 316 Buttino, I. See Ianora, A., 321
Buddemeier, R.W., 191 Button, D.K. See Druehl, L.D., 78
See Gattuso, J.P., 461 Buysse, D. See Beyst, B., 125
See Kleypas, J.A., 466 Buzas, M.A., 166
Buesseler, K.O. See Boyd, P.W., 456 See Culver, S.J., 167
See Hayek, L.C., 168
Buhl-Jensen, L., 126
Byers, J.E., 308
Bukantis, R.T. See Spencer, C.N., 136
Byrkjedal, I., 456
Bulboa, C.R., 308
Byrnes, J., 75
See Macchiavello, J., 325
Bull, G.D. See Babcock, R.C., 454
Bulleri, F., 396
See Benedetti-Cecchi, L., 395
C
See Moschella, P.S., 402 Cabrera, D. See Tovar, H., 340
Bullister, J.L. See McNeil, B.I., 469 Cáceres, C. See Gutierrez, A., 80, 319
Bunce, A., 456 Cáceres, H., 309
Bunge, J., 166 Cáceres, M., 309
Burau, J.R. See Kimmerer, W.J., 131 Caceres-Rubio, C.F. See Aguilar-Rosas, L.E., 74
Burbidge, A.A. See Ross, G.J.B., 472 Cadee, G.C. See Philippart, C.J.M., 470
Burch, T.L. See Osborne, A.R., 192 Cai, W., 456
Burd, F. See Cooper, N.J., 396 Caillaux, L. See Stotz, W., 338
Burford, M.A. See McKinnon, A.D., 468 Caillaux, L.M., 309
Burger, R.L. See Sandweiss, D.H., 334 Cailliau, C. See Garde, K., 461
Burgman, M.A., 75 Cairns, S. See Guinotte, J.M., 462
Burke, L., 396 Caldeira, K., 456
Burkett, V., 456 See Raven, J., 471
Burkholder, J.M. See Harvell, C.D., 463 Calderon, J. See Rutllant, J.A., 334
Burnett, W.J., 191 Calderon, R. See Fernández, M., 315
Burns, B.R. See Wigley, R.L., 138 Caldwell, M. See Madronich, S., 325
Burns, C.W. See Wilhelm, F.M., 138 Calliari, D., 126
Burns, F. See Beare, D.J., 454 See Rodríguez-Graña, L., 333
Burns, J.J. See Lowry, L.F., 132 Calvert, S.E. See Cowie, G.L., 312
Burrell, V.G., Jr. See Mackenzie, C.L., Jr, 401 Cambridge, M.L., 456
Burrows, M.T. See Emsley, S.M., 128 Camousseight, A. See Castro, S.A., 311
See Kendall, M.A., 465 Campbell, D. See Lyle, J.M., 468
485
AUTHOR INDEX
Campbell, R.A., 456 See Holbrook, S.J., 81

Campbell, S.J., 456 See Shanks, A.L., 336
See Baird, A.H., 190 Carr, M.R. See Olsgard, F., 170
Camus, P. See Thiel, M., 199–344 See Somerfield, P.J., 171
Camus, P.A., 75, 309 Carrasco, F. See Gutiérrez, D., 319
See Ahumada, R., 305 See Jaramillo, E., 322
See Castilla, J.C., 310 Carrasco, F.D., 309
See Vásquez, J.A., 87, 341 See Gallardo, V.A., 316
Canahuire, E. See Tarazona, J., 339 See Sepúlveda, R., 336
Canales, M.T. See Cubillos, L.A., 312 Carrasco, S., 309
Cancino, J. See Santelices, B., 335 Carreno, E. See Letelier, S., 324
Cancino, J.M. See Gajardo, G., 316 Carrette, T. See Little, M., 467
See Paine, R.T., 330 Carrington, E. See Helmuth, B., 463
See Sepúlveda, R., 336 Carriquiry, J.D. See Takesue, R.K., 339
Candia, M. See Ortlieb, L., 330 Carruthers, T.J.B., 457
Cane, M.A., 309 Carter, J.H.C., 126
Cañete, J.I. See Gallardo, V.A., 316 Carter, K. See Kleppel, G.S., 466
See Roa, R., 333 Cartes, C. See Palma, A.T., 331
Canfield, D.E. See Fossing, H., 316 See Vargas, C.A., 341
Cano, A. See Sandweiss, D.H., 334 Cartes, J.E., 126
Cano, J., 396 Cary, J. See Lemmens, J., 323
CansemiSoullard, M. See Delille, D., 313 Casanova, B. See De Jong-Moreau, L., 127
Cantillánez, M. See Avendaño, M., 306 Casanova, J.-P. See De Jong-Moreau, L., 127
Cantlon, J.E. See Odum, H.T., 170 Casas, G. See Orensanz, J.M., 134
Canty, P. See Ross, G.J.B., 472 Casas-Valdez, M.M. See Hernández-Carmona, G., 81
Cao, J.J. See Jickells, T.D., 465 Caselle, J.E. See Swearer, S.E., 339
Capella, J., 309 Casey, J. See O’Brien, C.M., 470
Capobianco, M. See Hamm, L., 399 Casotti, R. See Ianora, A., 321
Caputi, N., 456 Castel, J., 126
See Griffin, D.A., 462 Castiglioni, G.B. See Bondesan, M., 395
Caraco, N. See Duarte, C.M., 459 Castilla, A.C. See Prado, L., 332
Caragitsou, E. See Papaconstantinou, C., 134 Castilla, J.C., 76, 309, 310
Carbajal, G. See Rosenberg, R., 333 See Aiken, C., 305
Carcelles, A., 309 See Alvarado, J.L., 305
Card, M. See Hughes, T.P., 464 See Cerda, M., 311
Cárdenas, L., 309 See Clarke, M., 311
See Martínez, E.A., 326 See Correa, J.A., 312
See Poulin, E., 332 See Defeo, O., 313
Carder, K.L., 33 See Durán, L.R., 313
Caressa, S., 396 See Ebensperger, L.A., 313
Carlotti, F. See Bonnet, D., 455 See Fariña, J.M., 315
See Miller, C.B., 133 See Fernández, M., 315
Carlson, B.E. See Macke, A., 35 See Gelcich, S., 317
Carlsson, P., 456 See Jerardino, A., 322
Carlton, J.H., 126 See Kaplan, D.M., 322
Carney, R.S., 139–172, 166 See Kelaher, B.P., 322
Caro, A.U. See Castilla, J.C., 310 See Lagos, N.A., 323
Carola, M., 126 See Lardies, M.A., 323
See Coma, R., 126 See Manríquez, P.H., 325
Carotenuto, Y. See Ianora, A., 321 See Narváez, D.A., 329
Carpenter, R.C. See Stewart, H.L., 474 See Navarrete, S.A., 329
Carr, A., 457 See Paine, R.T., 330
See Mulkins, L.M., 133 See Piñones, A., 332
Carr, J.J. See Hannah, L., 399 See Poulin, E., 332
Carr, M.E., 309 See Roberts, C.M., 333
See Blanco, J.L., 307 See Sánchez, M., 334
See Thomas, A.C., 339 See Santelices, B., 335
Carr, M.H., 75, 76 See Sielfeld, W., 336
See Graham, M.H., 79 See Takesue, R.K., 339
486
AUTHOR INDEX
See Thiel, M., 199–344 Champalbert, G., 126

See Vásquez, J.A., 87, 341 Chan, B.K.K., 457
See Zacherl, D.C., 344 Chan, F. See Grantham, B.A., 318
Castillo, J.G., 310 See Leslie, H., 324
Castillo, M. See Aiken, C., 305 See Menge, B.A., 327
Castro, H. See Rodríguez, L., 333 Chan, H.R. See Webster, P.J., 476
Castro, L. See González, H.E., 317 Chang, H. See Boyd, P.W., 456
See Grünewald A., 318 Chang, J. See Colwell, R.K., 167
See Rodríguez-Graña, L., 333 Chang Chew Hung. See Hesp, P.A.,
Castro, L.R., 310, 311 Chansang, H., 191
See Hernández, E.H., 319 See Brown, B.E., 191
See Landaeta, M.F., 323 See Phongsuwan, N., 193
See Llanos-Rivera, A., 324 See Scoffin, T.P., 193
See Morales, C.E., 328 Chao, A., 166
See Pavez, M.A., 331 Chaplin, G. See Makris, N.C., 132
See Rodríguez-Graña, L., 333 Chapman, A.R.O., 76
See Thiel, M., 199–344 Chappell, J. See Wolanski, E., 476
See Vargas, C.A., 341 Chardy, P. See David, V., 127
See Yannicelli, B., 344 Charette, M. See Boyd, P.W., 456
Castro, S.A., 311 Chase, J.M. See Leibold, M.A., 323
Caswell, H. See Pineda, J., 170 Chavez, F. See Wang, H.J., 343
Catasti, V. See Yáñez, E., 344 Chavez, F.P., 311, 457
Caton, A., 457 See Montecino, V., 327
Catrilao, M. See González, J., 318 See Olivieri, R.A., 330
Cattaneo-Vietti, R. See Bavestrello, G., 394 Checkley, D.M. See Jahncke, J., 321
See Cicogna, F., 396 Cheeney, R.F. See Scoffin, T.P., 193
Cattrijsse, A. See Dewicke, A., 128 Chelazzi, L. See Colombini, I., 311
See Hampel, H., 129 Chemello, R. See Milazzo, M., 402
See Mees, J., 133 Chen, R. See Twilley, R.R., 475
Cauchy, A.L., 34 Chen, Y.C. See Lee, K.K., 467
Cauquil, E. See Hyder, P., 192 Cheng, D. See Harty, C., 463
Cavallini, F. See Simonini, R., 404 Cherr, G.N. See Garman, G.D., 79
Cayan, D.R. See McGowan, J.A., 468 468 See Huovinen, P.J., 81
See Venrick, E.L., 475 Chess, D.W., 126
Ceballos, S. See Bonnet, D., 455 Chess, J.R. See Hobson, E.S., 81
Cebrián, J. See Duarte, C.M., 459 Chia, F.S. See Hurd, C.L., 81
See Hauxwell, J., 399 Chiaverini, D. See Meinesz, A., 401
See Marbà, N., 401 Childers, D.L. See Beck, M.W., 394
Ceccherelli, G. 396 Chin, N.K.M., 76
See Milazzo, M., 402 See Brown, M.T., 75
Ceccherelli, V.U. See Martin, D., 401 Chipman, D. See Langdon, C., 466
Cech, J.J., Jr. See Roessig, J.M., 472 Chisholm, J.R.M. See Jaubert, J.M., 400
Cencini, C., 396 Chisholm, S.W. See Moore, L.R., 469
Cerda, G. See Sielfeld, W., 336 Chisolm P. See Kaufman L., 322
Cerda, M., 311 Chmielewski, F.M. See Menzel, A., 469
See Castilla, J.C., 310 Choi, J.S., 126
Cerrano, C. See Bavestrello, G., 394 See Frank, K.T., 129
Cervetto, G. See Calliari, D., 126 Chong, J., 311
Ceschia, C. See Caressa, S., 396 Choo, K.S., 457
Chacana, M. See Westermeier, R., 343 Chopin, T., 311
Chadwick, D.B. See Tegner, M.J., 87 Choquet, R. See Rivalan, P., 472
Chaigneau, A., 311 Chou, L. See Engel, A., 460
Chakraborty, P.P. See Khan, P.K., 192 Chou Loke Ming. See Hesp, P.A.,
See Pal, T., 192 Chouinard, G.A. See Hanson, J.M., 130
Chaloupka, M., 457 Christensen, J.P. See Codispoti, L.A., 311
Chamberlain, Y.M., 76 Christianen, M.J.A. See Dorenbosch, M., 459
Chambers, D. See Allan, R., 190 Christie, H., 396
Chambers, L.E., 457 Chu, D. See Lawson, G.L., 132
Chamot-Rooke, N. See Pubellier, M., 193 Chubb, C. See Caputi, N., 456
487
AUTHOR INDEX
Chubb, C.F. See Griffin, D.A., 462 Coles, R.G. See Preen, A.R., 471
Chumán de Flores, E. See Rosenberg, R., 333 Coley, T.L. See Coale, K.H., 311
Church, J. See Walsh, K.J.E., 476 Collantes, G., 311
Church, J.A., 457 Collier, M.A. See Gordon, H.B., 462
See Gregory, J.M., 462 Collins, A.G. See Castilla, J.C., 310
Chýlek, P., 34 Collins, J.D. See Druehl, L.D., 77
Cicogna, F., 396 Colman, J.S., 126
See Russo, G.F., 403 See Russell, F.S., 472
Cid, L. See Acuña, E., 304 Coloma, C., 311
Cifuentes, M. See Fernández, M., 315, 316 See Marchant, M., 325
Cifuentes, S. See Brante, A., 308 Colombini, I., 311
Cimino, J. See Pruett, L., 403 Colucci-D’Amato, L. See Ianora, A., 321
Cinelli, F. See Benedetti-Cecchi, L., 395 Colwell, R.K., 167
See Boudouresque, C.F., 395 See Gotelli, N.J., 168
See Ceccherelli, G. 396 Colwell, R.R. See Harvell, C.D., 463
See Meinesz, A., 401 Coma, R., 126
See Piazzi, L., 403 See Carola, M., 126
Cirik, S. See Boudouresque, C.F., 395 See Ribes, M., 134
Cisneros-Mata M.A. See Porteiro C., 471 Compagna, C. See Boersma, D., 307
Clabaut P. See Pasqualini V., 402 CONAMA-PNUD, 311
Clapin, G. See Lemmens, J., 323 Condie, S.A. See Ridgway, K.R., 472
Clark, R.P., 76 Conkright, M.E., 311
Clarke, A., 166, 311 Conn, J. See Hutchins, D.A., 321
See Rex, M.A., 170
Connell, J.H. See Hughes, T.P., 464
Clarke, K.R., 166, 191
See Tutschulte, T.C., 87
See Ellingsen, K.E., 167
Connell, S.D., 396
See Warwick, R.M., 172
See Fowler-Walker, M.J., 460
Clarke, M., 311
See Gorgula, S.K., 462
Connelly, X.M. See Detres, Y., 458
Clarke, P.J., 457
Connolly, R.M. See Loneragan, N.R., 467
Clarke, R.B., 166
See Seddon, S., 473
Clarke, W.D., 76
Connolly, S.R., 167, 311
See Rosenthal, R.J., 85
See Hughes, T.P., 464
Clavano, W.R., 1–38
Connor, D.W., 396
Clayton, M.N. See Wiencke, C., 476
Conover, R.J., 127
Clement, A. See Sandweiss, D.H., 334
Clements, F.E., 166 Conte, M. See Antia, A.N., 454
Clementson, L. See Harris, G., 463 Contreras, H., 312
Clementson, L.A. See Harris, G.P., 463 See Duarte, C., 313
See Thresher, R.E., 474 See Dugan, J.E., 313
Clendenning, K.A. See Wing, B.L., 88 See Jaramillo, E., 322
Closs, G.P. See Sutherland, D.L., 136 See Quijón, P., 332
Clough, J. See Galbraith, H., 461 Contreras, M. See Simeone, A., 337
Clutter, R.I., 126 Convey, E. See Walther, G.R., 476
See Fleminger, A., 128 Cook, K. See Bonnet, D., 455
Coale, K.H., 311 Cooke, R. See Jackson, J.B.C., 399
Coates, M., 457 Cooke, R.G. See Lotze, H.K., 401
Cocito, S. See Sandulli, R., 403 See Pandolfi, J.M., 402
Coddington, J.A. See Colwell, R.K., 167 Cooper, N.J., 396
Codignotto, J.O. See Burkett, V., 456 Cooper, T. See Cooper, N.J., 396
Codispoti, L.A., 311 Corbert, A.S. See Fisher, A.A., 167
Coelho, S.M. See Rijstenbil, J.W., 472 Corbett, D. See Steneck, R.S., 86
Cohen, C.S., 311 See Steneck, R.S., 404, 474
Cohen, P.J., 126 Cordell, J. See Dean, A.F., 127
Colangelo, M.A. See Martin, D., 401 Cordi, B., 457
Cole, D.W. See Henry, E.C., 80 Córdova, C. See Fernández, E., 315
Cole, J. See Wang, H.J., 343 Cordova, J. See Bertrand, A., 307
Cole, R.G., 457 Corkeron, P.J. See Marsh, H., 468
Coleman, F.C., 457 Corliss, L.A. See Mrosovsky, N., 469
488
AUTHOR INDEX
Cormaci, M., 396 Crowley, G.M. See Garnett, S.T., 461

See Boudouresque, C.F., 395 Croxall, J.P. See Phillips, R.A., 332
Cornejo, M., 312 Crozier, R.H., 167
Cornejorodriguez, M.D. See Enfield, D.B., 314 Cruz-Palacios, V., 457
Cornelius, J. See King, B.R., 466 CSIRO, 457
Cornet, M., 127 Cubasch, U., 458
Cornuelle, B. See Willis, J.K., 476 Cubillos, L. See Ibáñez, C.M., 321
Correa, J.A., 312 See Neira, S., 330
See Neill, P.E., 329 Cubillos, L.A., 312
See Faugeron, S., 315 See Arancibia, H., 305
See Santelices, B., 335 See Arcos D.F., 305
Correa, T., See Gutierrez, A., 80, 319 Cuesta, J.A. See Drake, P., 128
Cortés, M. See Thiel, M., 199–344 Cuevas, L.A., 312
Costanza, R., 396, 457 See Troncoso, V.A., 340
Costello, M.J., 167 See Vargas, C.A., 341
Côté, I.M. See Fish, M.R., 460 Culik, B.M. See Hennicke, J.C., 319
See Gardner, T.A., 461 See Luna-Jorquera, G., 325
Cottalorda, J.-M. See Meinesz, A., 401 Cullen, J.J. See Kouwenberg, J.H.M., 466
Cottenie, K. See Halpern, B.S., 80 Culver, S.J., 167
Coulon, A.R. See Anderson, B.S., 74 See Buzas, M.A., 166
Courtenay, S.C. See St-Hilaire, A., 136 Cummings, V. See Thrush, S.F., 474
Coury, D.A. See Reed, D.C., 85 Cummings, V.J. See Lohrer, A.M., 467
Cowan, T. See Cai, W., 456 See Norkko, A., 470
Cowen, R.K., 76, 457 See Thrush, S.F., 171, 475
Cowie, G.L., 312 See Turner, S.J., 404
Cowie, R.J. See Both, C., 455 Cunha, M.R., 127
Cowles, R.P., 127 Cunningham, A. See MacCallum, I., 35
Coyer, J.A., 76 Cunningham, C.W. See Wares, J.P., 343
See Holbrook, S.J., 81 Cuq, F. See Glemaréc, M., 398
Coyne, G. See Rex, M.A., 170 Curiel, D. See Rismondo, A., 403
Craig, E.A. See Becker, J., 307 Curnel, Y. See Menzel, A., 469
Craig, M.G. See Polovina, J.J., 471 Curray, J.R., 191
Craigie, J.S. See Chapman, A.R.O., 76 Currie, B. See Weeks, S.J., 343
Crain, J.A., 127 Currie, V. See Dayton, P.K., 76
Crame, J.A. See Clarke, A., 166 See Dayton, P.K., 313
See Rex, M.A., 170 Curry, J.A. See Webster, P.J., 476
Crandall, W.C., 76 Curtis, T.G.F., 396
Crater, T.S. See Bickmore, B.R., 33 Cury, P., 312
Crawford, R.J.M. See Cury, P., 312 Cushing, D.H., 312, 458
Crecelius, E.A. See Huesemann, M.H., 464
Creese, R.G. See Cole, R.G., 457
Crepinsek, Z. See Menzel, A., 469 D
Cresswell, G. See Blackburn, S.I., 455
Cresswell, G.R. See Maxwell, J.G.H., 468 Daborn, G.R. See Stone, H.H., 136
Cribb, A.B., 76 Dadswell, M.J. See Carter, J.H.C., 126
Crick, H.Q.P., 457 da Fonseca, G.A.B. See Brooks, T.M., 395
See Learmonth, J.A., 466 Dahl, E. See Brattstöm, H., 308
See Robinson, R.A., 472 Dahlberg, K. See Jansson, B.-O., 400
Crisci, J.V. See Posadas, P., 170 Dahlgren, C.P. See Harborne, A.R., 463
Crisp, D.J., 312 See Sobarzo, M., 337
Crist, T.O., 167 Dahlhoff, E., 312
See Veech, J.A., 172 Dahlin, H. See Rosenberg, R., 403
Cronin, T.W. See Losey, G.S., 467 Dalh, A. See Menzel, A., 469
Croot, P. See Boyd, P.W., 456 Dall, W. See Staples, D.J., 474
Crosa, G. See Marchetti, R., 401 Dall, W.H., 312
Crosbie, N.D. See Furnas, M.J., 461 Dam, H. See Peterson, W.T., 331
Crossley, A.C. See Sedwick, P.N., 473 D’Amato, A.F. See Godfrey, M.H., 462
Crowder, L.B. See Heck, K.L., Jr, 399 Damkaer, D.M., 458
Crowe, T.P. See Thompson, R.C., 404 See Williams, A.B., 138
489
AUTHOR INDEX
Daneri, G., 312 Dean, T.A., 77

See Cuevas, L.A., 312 See Deysher, L.E., 77, 458
See Escribano, R., 314 See Schroeter, S.C., 86
See González, H.E., 317 See Tegner, M.J., 87
See Troncoso, V.A., 340 DeAngelis, D.L. See Grimm, V., 129
See Vargas, C.A., 341 De’ath, G. See Brodie, J., 456
Danielsson, K. See Rudstam, L.G., 135 See De Vantier, L.M., 458
Dann, P. See Ross, G.J.B., 472 See McKinnon, A.D., 468
D’Anna, G. See Badalamenti, F., 394 Debinski, D.M. See Su, J.C., 171
Dantagnan, P., 312 Defeo, O., 313
d’Arge, R. See Costanza, R., 396, 457 See Castilla, J.C., 310
Darling, J.D., 127 See Contreras, H., 312
Darnaude, A.M. See Salen-Picard, C., 334 Defile, C. See Menzel, A., 469
Darrigran, G. See Orensanz, J.M., 134 deFrance, S.D. See Keefer, D.K., 322
Darwin, C., 191 DeGange, A.R., 127
Darwin, C.R., 76, 312 De Grave, S., 396
Dashfield, S.L. See Warwick, R.M., 172 de Groot, R. See Costanza, R., 396, 457
Da Silva, A.M., 312 de Groot, S.J. See Lindeboom, H.J., 401
Dauby, P.A., 127 de Gruijl, F.R. See Kelfkens, G.,
Dauvin, J.-C., 127 de Gusmao, L.L.F. See Gama, A.M. da S., 129
See Elizalde, M., 128 de Haan, J.F. See Volten, H., 36
See Mouny, P., 133 Deibel, D. See Richoux, N.B., 135
See Vallet, C., 137 de Jager, D., 458
See Wang, Z., 137 De Jong-Moreau, L., 127
See Zouhiri, S., 138 Dekhuijzen, A.J. See Both, C., 455
Dauwe, B., 127 Dekker, R. See Philippart, C.J.M., 470
Davenport, J., 458 de la Huz, Ch. See Barbera, C., 394
Davenport, J.L. See Davenport, J., 458 DeLancey, L.B., 127
David, P.M., 127 de Lange, H..J. See Hessen, D., 463
David, V., 127 de Lange, P.J. See de Lange, W.P., 458
Davidson, N.C., 396 de Lange, W.P., 458
Davies, C.E., 396 Delaune, R.D.,See Parkinson, R.W., 470
Davies, P. See Babcock, R., 454 Delépine, R., 77
See Harris, G.P., 463 See Asensi, A., 74
Davis, A. See Wahl, M., 475 Delgado, L., 127
Davis, A.R., 458 Delgado, L.E. See Marín, V.H, 326
See Przeslawski, R., 471 Delgado, O., 396
Davis, C.S. See Kleppel, G.S., 466 Delille, B., 77
Davis, G., 458 See Engel, A., 460
Davis, R.F. See Kouwenberg, J.H.M., 466 Delille, D., 313
Davison, D.M., 396 See Delille, B., 77
Dawson, E.Y., 76 Dell, R.K., 313
Dawson, S.P., 458 Della Croce, N. See Thurston, M.H., 340
Day, R.W. See Zacherl, D.C., 344 Dellarossa, V. See Daneri, G., 312
Day-Lewis, A. See Keefer, D.K., 322 Dellinger, T., 313
Dayton, P.K., 76, 77, 313, 396, 458 DeMartini, E.E., 77
See Barilotti, D.C., 75 See Plummer, K.M., 134
See Genin, A., 461 See Polovina, J.J., 471
See Graham, M.H., 79 DeMaster, D.J. See Smith, C.R., 337
See Kinlan, B.P., 82 DeMaster, D.P. See Laidre, K.L., 82
See Rosenthal, R.J., 85 Demetropulous, A. See Jeftic, L., 400
See Sala, E., 334 Den Hartog, C., 397
See Seymour, R.J., 86 Dennison, W. See Walker, D., 475
See Tegner, M.J., 86, 87, 474 Dennison, W.C. See Abal, E.G., 453
See Thrush, S.F., 404 See Carruthers, T.J.B., 457
See Vetter, E.W., 342 See Dawson, S.P., 458
See Vilchis, L.I., 475 Denny, M., 458
De Amesti, P. See Stotz, W., 338 Denny, M.W. See Li, N.K., 467
Dean, A.F., 127 See Utter, B.D., 87
490
AUTHOR INDEX
De Oliveira J.A.A. See Porteiro C., 471 Dixon, J.D. See Dean, T.A., 77
Department of Marine and Coastal Resources, Thailand, See Schroeter, S.C., 86
191 Dixon, K.W. See Gregory, J.M., 462
de Perera, T.B. See Garcia, C.M., 461 Djellouli, A. See Meinesz, A., 401
Depledge, M.H. See Cordi, B., 457 Djurfeldt, L. See Shaffer, G., 336
De Pol, R. See Ulloa, O., 340 Dobow, J. See Wilcove, D.S., 405
De Pol-Holz, R. See Atkinson, L.P., 306 Dobretsov, S. See Wahl, M., 475
Deprez, T., 128 Dobson, A.P. See Harvell, C.D., 463
Dermott, R.M. See Johannsson, O.E., 130 Dodd, J., 397
De Robertis, A., 128 Dodson, J.J. See Winkler, G., 138
Desai, B.N., 191 Doerries, M.B., 167
Desalle, R. See Herbst, L., 463 Döhler, G., 458
Descimon, H. See Parmesan, C., 470 Dolman, P.M. See Sutherland, W.J., 338
Desprez, M., 397 Domin, A. See Wilhelm, C., 476
Desqueyroux, R., 313 Donahoe, C.J., 128
Desqueyroux-Faúndez, R. See Castilla, J.C., 310 Donders, T.H. See Sangiorgi, F., 403
Destombe, C. See Faugeron, S., 315 Done, T.J., 459
Detres, Y., 458 See De Vantier, L.M., 458
Dette, H.H. See Hamm, L., 399 See Orr, J.C., 470
Devinny, J.S., 77 Doney, S.C. See Boyd, P.W., 456
Devlin, M.J., 458 See Sarmiento, J.L., 473
DeVried, P.J. See Lande, R., 169 Donkin, M.E. See Cordi, B., 457
De Vantier, L.M., 458 Donnan, D.W., 397
de Vaugelas, J. See Meinesz, A., 401
Donnellan, M.D., 77
de Villèle, X., 397
Donnelly, A. See Menzel, A., 469
de Vries, M. See Martin, D., 401
Donner, S.D., 459
Dewees, C.M. See Leet W.S., 82
Doody, J.P., 397
Dewey, R. See Kunze, E., 131
See Davidson, N.C., 396
Dewicke, A., 128
Dorenbosch, M., 459
See Beyst, B., 125
Dorman, L.M. See McGowan, J.A., 468
Dewitte, B. See Ramos, M., 333
Dortch, Q. See Thessen, A.E., 474
DeWreede, R.E., 77
dos Santos, A. See Bonnet, D., 455
Dey, D.B. See Damkaer, D.M., 458
Doumenge, F. See Barnabe, G., 394
Deysher, L. See Grove, R.S., 462
Dower, J.F. See Kunze, E., 131
Deysher, L.E., 77, 458
Downes, B.J., 77
See Tegner, M.J., 87
Dezileau, L. See Muñoz, P., 329 See Keough, M.J., 322
Díaz, A. See Palma, A.T., 331 Downing, K. See Boyd, P.W., 456
Díaz, C. See Acuña, E., 304 Downs, C.A., 191
Diaz, H.F., 77 See Brown, B.E., 191
Diaz, R.J., 397 Drake, C.M. See Davidson, N.C., 396
Diaz-Almela, E., 458 Drake, P., 128
DiBacco, C., 313 Drake, P.T. See Reed, D.C., 85
Diebold, E.N. See Simeone, A., 337 Drew, E.A., 191
Dieckmann, U., 128 Dring, M.J. See Birkett, D.A., 395
Diehl, S., 458 Drinkwater, K. See Choi, J.S., 126
Diekman, R. See Bonnet, D., 455 Drinkwater, K.F., 459
Dijkema, K.S., 397 Drohan, A.F., 459
Dimmlich, W.F. See Ward, T.M., 476 Drosdowsky, W. See Allan, R., 190
Dinesen, G.E. See Moschella, P.S., 402 Druehl, L.D., 77, 78
Dippner, J.W. See Kroencke, I., 323 See Lane, C.E., 82
Dittmann, S. See Grimm, V., 129 See Saunders, G.W., 85
DiTulliio, G.R. See Sedwick, P.N., 473 See Swanson, A.K., 86, 474
See Hutchins, D.A., 321 See Wheeler, W.N., 88
See Tortell, P.D., 475 Du, T. See Gordon, H.R., 34
Divoky, G.J., 128 Duarte, C., 313
Dix, M. See Cubasch, U., 458 See Dugan, J.E., 313
Dix, M.R. See Gordon, H.B., 462 See Jaramillo, E., 322
Dixon, J., 77 Duarte, C.M., 397, 459
491
AUTHOR INDEX
See Diaz-Almela, E., 458 See Hempel, W.M., 80

See Hemminga, M.A., 399 See Reed, D.C., 85, 333
See Marbà, N., 401 Ebensperger, L.A., 78, 313
Duarte, W.E., 313 See Villegas, M.J., 343
See Moreno, C.A., 328 EC, 397
See Zamorano, J.H., 344 Eckerle, L. See Brante, A., 308
Dubinsky, Z. See Stambler, N., 194 Eckert, S.A., 459
Dubovik, O., 34 Edding, M., 313, 314
See Zhao, T.X.-P., 36 See Macchiavello, J., 325
Duce, R.A. See Jickells, T.D., 465 See Tala, F., 339
Duchene, J.C. See Guizien, K., 318 See Véliz, K., 342
Ducklow, H.W., 313 Edgar, G. See Walker, D., 475
Dudley, C. See Crick, H.Q.P., 457 Edgar, G.J., 78, 397, 459
Duff, K.L. See Davidson, N.C., 396 Edmonds, S.J. See Womersley, H.B.S., 477
Duffus, D.A. See Dunham, J.S., 128 Edmunds, P.J. See Gleason, D.F., 461
Duffy, D., 313 Eduardo, C., 167
Dufresne, J.L. See Bopp, L., 455 Edvardsen, E. See Jonzén, N.,
Dugan, J. See Roberts, C.M., 333 Edwards, A.J. See Mumby, P.J., 469
Dugan, J.E., 313 Edwards, M., 459
See Airamè, S., 393 See Beaugrand, G., 454
See Bomkamp, R.E., 307 See Bonnet, D., 455
Dugan, P., 397 See Kirby, R.R., 466
Dugdale, R.C. See Hogue, V.E., 464 Edwards, M.J., 459
Duggan, S. See McKinnon, A.D., 468 Edwards, M.S., 78, 314, 459
Duggins, D.O., 78 See Clark, R.P., 76
See Estes, J.A., 78 Edwards, P.B. See Dayton, P.K., 77, 396, 458
Duke, N.C., 459 See Tegner, M.J., 87, 474
Dukowicz, J.K. See Griffies, S.M., 462 Edwards-Jones, G. See Gelcich, S., 317
Dulvy, N.K., 459 Edyvane, K.S., 459
Dumont, C.P. See Thiel, M., 199–344 See Seddon, S., 473
Dunham, J.S., 128 EEA., 397
Dunlop, J.N., 459 Eekhout, S. See Bustamante, R.H., 308
Dunn, J.R. See Ridgway, K.R., 472 Eeva, T. See Both, C., 455
Dunne, J.A., 128 Eggiman, D.W. See Carder, K.L., 33
Dunne, R.P., 191 Eggleston, D.B. See Beck, M.W., 394
See Brown, B.E., 191 Ego, F. See Pubellier, M., 193
Dunton, K., 313 Ehlers, A. See Lotze, H.K., 401
Dunton, K.H., 78 Ehrhart, L.M. See Weishampel, J.F., 476
Dupont, F. See St-Hilaire, A., 136 Eijsackers, M. See Rijstenbil, J.W., 472
Durán, L.R., 313 Eilers, M.R. See Matilla, J., 132
See Castilla, J.C., 310 Eischeid, J.K. See Diaz, H.F., 77
Durán, R. See Sielfeld, W., 336 Eissler, Y., 314
Durante, K.M., See Hurd, C.L., 81 Ekebom, J., 398
Durr, S.T. See Wahl, M., 475 Ekman, S., 314
Dürselen, C.-D. See Hillebrand, H., 34 El Abed, A. See Meinesz, A., 401
Dutton, P.H. See Polovina, J.J., 471 Elderfield, H. See Raven, J., 471
Duysens, L.M.N., 34 Eleftheriou, A., 128
Elías, R. See Orensanz, J.M., 134
Elizalde, M., 128
E Elkins, J.W. See Codispoti, L.A., 311
Ellenberg, U., 314
Eaglesham, G. See Negri, A., 469 See Mattern, T., 326
Eakin, C.M. See Kleypas, J.A., 466 See Simeone, A., 337
Eardley, D.D., 78 Ellingsen, K.E., 167
See Hempel, W.M., 80 See Ugland, K. I., 172
Ebeling, A.W., 78 Elliott, T.I. See Gordon, H.B., 462
See Bray, R.N., 75 Ellis, D., 167
See Eardley, D.D., 78 Ellis, J.C., 314
See Harris, L.G., 80 See Fariña, J.M., 315
492
AUTHOR INDEX
Ellis, J.I. See Norkko, A., 470 Espíndola, F. See Yáñez, E., 344
See Thrush, S.F., 474 475 Espinoza, F., 315
Ellis, J.R. See Perry, A.L., 470 Espinoza, R. See Buschmann, A.H., 75
Ellison, A.M., 459 See Buschmann, A.H., 308
See Farnsworth, E.J., 460 Espinoza, S. See Marín, V.H., 326
Ellison, J.C., 460 Essink, K. See van Dalfsen, J. A., 405
Elmgren, R. See Hansson, S., 130 Estes, J. See Steneck, R.S., 86
See Rosenberg, R., 403 Estes, J.A., 78
Elmi, C. See Bondesan, M., 395 See Duggins, D.O., 78
El-Sayed, S., 459 See Erlandson, J.M., 78
Emanuel, K., 460 See Jackson, J.B.C., 399
Emblow, C.S. See Costello, M.J., 167 See Paddack, M.J., 84
Emes, C. See Speirs, D., 136 See Steneck, R.S., 474
Emmerson, M., 128 See Steneck, R.S., 404
Emsley, S.M., 128 Estes, J.E. See Jensen, J.R., 81
Enbysk, B.J., 128 Estrella, N. See Menzel, A., 469
Endean, R., 460 Etter, R.J. See Levin, L.A., 169, 324
Ene, A., 460 See Rex, M.A., 170
Ene, A. See Herbst, L., 463 See Witman, J.D., 172
Enemar, A. See Both, C., 455 EUCC — The Coastal Union, 398
Enfield, D.B., 314 Evans, B.T.N. See Fournier, G.R., 34
Engdhal E.R. See Bilham, R., 191 Evans, M.J., 460
Engel, A., 460 Evans, M.S., 128
Engkvist, R. See Nilsson, J., 402 Ewel, K.C., 460
See Worm, B., 405 Ezzi, I.A. See Gibson, R.N., 129
Enríquez, S. See Marbà, N., 401
Enríquez-Briones, S. See Gallardo, V.A., 316
See Roa, R., 333 F
Environment Australia, 460
Eppley, R.W., 460 Faaborg, J. See Terborgh, J.W., 171
Epstein, P.R. See Harvell, C.D., 463 Fabbri, D. See Trombini, C., 404
Erftemeijer, P.L.A. See Ochieng, C.A., 470 Fabri, M.C., 167
Erga, S.R., 460 Fabricius, K. See Brodie, J., 456
Eriksson, B.K., 398 See Negri, A., 469
See Johansson, G., 400 Fabricius, K.E., 460
Erkkilä, A. See Ekebom, J., 398 See De Vantier, L.M., 458
Erlandson, J. See Jackson, J.B.C., 399 Fabry, V.J. See Orr, J.C., 470
Erlandson, J.M., 78 Fadii, N. See Baird, A.H., 190
See Graham, M.H., 79 Fagan, W.F. See Andelman, S.J., 166
See Kinlan, B.P., 82 Fahrbach, E. See Arntz, W., 306
See Steneck, R.S., 86, 404, 474 Fain, S.R., 78
Ernst, B. See Roa, R., 333 Faith, D.P., 167
Escribano, N. See Marín, V.H., 326 FAL, 315
See Ulloa, O., 340 Fallaci, M. See Colombini, I., 311
Escribano, R., 314, 315 Fanelli, G., 398
See Castro, L.R., 311 FAO, 315
See Fernández, D., 315 Farber, S. See Costanza, R., 457
See Giraldo, A., 317 See Costanza, R., 396
See González, H.E., 317, 318 Farías, L., 315
See Herrera, L., 319 See Cornejo, M., 312
See Hidalgo, P., 320 See Escribano, R., 314
See Ortlieb, L., 330 See Graco, M., 318
See Palma, W., 331 Farías, M. See Fonseca, T., 316
See Rendell, L., 333 Fariña, J.M., 315
See Rodríguez, L., 333 See Ellis, J.C., 314
See Rojas, P.M., 333 See Sabat, P., 334
See Takesue, R.K., 339 Farnsworth, E.J., 460
See Thiel, M., 199–344 See Ellison, A.M., 459
See Vargas, C.A., 341 Farrell, T.M., 315
493
AUTHOR INDEX
Faugeron, S., 315 Fishelson, L. See Moran, S., 133

See Neill, P.E., 329 Fisher, A.A., 167
Fauth, J.E. See Downs, C.A., 191 Fisher, N.S. See Twining, B.S., 137
Feder, H.M., 78 Fitt, W.K. See LaJeunesse, T.C., 192
Feely, R.A. See Orr, J.C., 470 See Wellington, G.M., 476
Fehner, U. See Antia, A.N., 454 Fitzpatrick, L. See Guerra, C., 318
Feld, N. See Bilham, R., 191 Fitzpatrick, M. See Bunge, J., 166
Feldman, M.W. See Zhivotovsky, L.A., 138 Fitzwater, S.E. See Coale, K.H., 311
Félix-Pico, E. See Narvarte, M.E., 329 See Martin, J.H., 326
Félix-Uraga R. See Porteiro C., 471 Flato, G.M. See Gregory, J.M., 462
Feller, R.J. See Zagursky, G., 138 Fleischer, S. See Rosenberg, R., 403
Fengshan, X. See Rhoads, D.C., 471 Fleminger, A.,128
Fenner, D. See Turner, J.R., 194 See Marín, V.H., 326
Ferl, R.J. See Bowen, B.W., 455 Flensted-Jensen, E. See Moller, A.P., 469
Fernández, D., 315 Fletcher, T.D. See Keatley, M.R., 465
Fernández, E., 315 Flick, P. See Brooks, T.M., 395
See Lleellish, J., 324 Flinkman, J., 128
See Lleellish, M., 83 See Viherluoto, M., 137
Fernández, M., 315, 316 See Viitasalo, M., 137
See Astorga, A., 306 Flint, E.N. See Polovina, J.J., 471
See Baeza, J.A., 306 Flores, C. See Pizarro, J., 332
See Brante, A., 308 Flores, L.A. See Rosenberg, R., 333
See Brokordt, K.B., 308 Flores, M. See Bernal, M., 307
Flores, V. See Santelices, B., 335
See Escribano, R., 314
Flores, V.A. See Arntz, W., 306
See Marquet, P.A., 326
Flynn, A.J., 128
See Rivadeneira, M.M., 333, 472
Flynn, M.N. See Tararam, A.S., 137
See Ruiz-Tagle, N., 334
Foertch, J.F. See Keser, M., 466
Fogarty, M.J., 129
Fernandez de Puelles, M.L. See Bonnet, D., 455
Foggo, A., 167
Fernández-Delgado, C. See Drake, P., 128
Foley, A.M., 460
Fernex, F. See Jeftic, L., 400
Foley, M.M. See Menge, B.A., 327
Ferraroli, S., 460
Folke, C. See Hughes, T.P., 464
Ferreyra, G.A. See Hernando, M.P., 463
Follegati, R. See Ortlieb, L., 330
Fetter, M. See Möllmann, C., 133
Fonck, E. See Santelices, B., 335
Fey, C. See Haury, L., 130
Feyrer, F., 128 Fonck, E.A. See Edding, M., 313, 314
Fiala, M. See Delille, B., 77 See Vásquez, J.A., 341
Fick-Child, K.J. See Foley, A.M., 460 Fong, P. See Smith, J.R., 473
Fiedler, P.C., 316 Fonseca, C.R., 460
Field, C.D., 460 Fonseca, M.S., 460
Fielding, P.J. See Attwood, C.G., 74 Fonseca, T., 316
Figueroa, C. See Takesue, R.K., 339 See Johnson, D.R., 322
Figueroa, D., 316 Forbes, D.L. See Burkett, V., 456
See Atkinson, L.P., 306 Forbes, E., 167
See González, H.E., 317, 318 Ford, E., 167
See Rutllant, J.A., 334 Ford, R. See Lawrie, S.M., 132
See Sobarzo, M., 337 Fordham, E. See Frid, C.L.J., 460
See Yannicelli, B., 344 Forster, S. See Fossing, H., 316
Figueroa, F.L. See Gómez, I., 317 Fortuna, S. See Wilkie, M.L., 405
See Huovinen, P., 321 Forward, R.B., 316
Filella, Y. See Menzel, A., 469 See Goddard, S.M., 129
Filún, L. See Buschmann, A.H., 75, 308 Fosberg, F.R., 78
Finger, I. See Rosenberg, R., 333 Fosså, J.H., 129
Finke, G.R. See Navarrete, S.A., 329 See Attramadai, Y.G., 125
Finlayson, C.M., 398 See Buhl-Jensen, L., 126
Firme, G.F. See Hutchins, D.A., 321 Fossing, H., 316
Fischer, G. See Antia, A.N., 454 Foster, E.G., 129
Fish, M.R., 460 Foster, M.S. 78, 79
494
AUTHOR INDEX
See McConnico, L., 83 Frost, M.T. See Foggo, A., 167

See Clark, R.P., 76 Fry, B. See Hansson, S., 130
See Cowen, R.K., 76 Fry, E.S. See Voss, K.J., 36
See Graham, M.H., 79 Fryd, M. See Nielsen, T.G., 192
See Hernández-Carmona, G., 80 Fuchs, B.M. See Kuypers, M.M., 323
See Kimura, R.S., 82 Fuenteseca, J. See Marín, V.H., 326
See Reed, D.C., 85, 471 Fuenzalida, H. See Rutllant, J.A., 334
See Schiel, D.R., 86, 473 Fuenzalida, R., 316
See Spalding, H., 86, 474 Fuller, P.J. See Ross, G.J.B., 472
Fournier, G. See Jonasz, M., 34 Fulton, E.A., 461
Fournier, G.F. See Jonasz, M., 34 Fulton, R.S., III, 129
Fournier, G.R., 34 Funnell, G. See Turner, S.J., 404
Fourqurean, J.W., 460 460 Funnell, G.A. See Norkko, A., 470
Fowler-Walker, M.J., 460 See Thrush, S.F., 475
Fox, C.J. See O’Brien, C.M., 470 Furlani, D.M. See Thresher, R.E.,
Fox, D.S. See Grantham, B.A., 318 Furlong, C. See Hauxwell, J., 399
Fox, G. See Ball, B.J., 394 Furnari, G. See Cormaci, M., 396
Fox, G.E. See Woese, C.R., 344 Furnas, M.J., 461
Fox, R. See Hickling, R., 463 See McKinnon, A.D., 468
Francis, C.M. See Marra, P.P., 468
Furness, R., 316
Francis, R.C. See Mantua, N.J., 468
Furukawa, K., 461
Franco, E. See Ariani, A.P., 125
Furusawa, M. See Amakasu, K., 125
Francour P. See Perez, T., 470
Frank, K. See Gray, J.S., 168
Frank, K.T., 129
See Choi, J.S., 126
G
Franke, H.D., 398 Gabbbianelli, G. See Bondesan, M., 395
Frankignoulle, M. See Delille, B., 77 Gabrielides, G.P. See Jeftic, L., 400
See Delille, D., 313 Gabrielson, P.W., 79
See Gattuso, J.P., 461 Gacia, E. See Moschella, P.S., 402
Fraschetti, S. See Guidetti, P., 398 Gage, J., 167
Frazier, S. See Nivet, C., 402
See Levin, L.A., 324
Frederich, M., 316
See Narayanaswamy, B.E., 170
Fredriksen, S., 79
See Roberts, J.M., 472
See Christie, H., 396
Gaines, S. See Roughgarden, J., 85, 333
Freeland, H.J., 316
Gaines, S.D. See Broitman, B.R., 308
Freidenburg, T.L. See Menge, B.A., 327
See Gaylord, B., 461
Freiwald, A. See Guinotte, J.M., 462
See Kinlan, B.P., 322
See Roberts, J.M., 472
See Lubchenco J., 169
Frere, E., 316
See Menge, B.A., 327
Frette, O. See Erga, S.R., 460
Freudenthal, T., 319 See Siegel, D.A., 336
Frew, R. See Boyd, P.W., 456 See Wares, J.P., 343
Frid, C., 398 See Zacherl, D., 477
Frid, C.L., 316 See Zacherl, D.C., 344
Frid, C.L.J., 398, 460 See Blanchette, C.A., 75
Friedel, M.H., 461 Gajardo, G., 316
Friederich, G.E. See Codispoti, L.A., 311 Gajardo, J.A. See Thiel, 199–344
Friedman, C.S. See Vilchis, L.I., 475 Gal, G., 129
Friedman, J.D. See Vilchis, L.I., 475 See Johannsson, O.E., 130
Friesen, J.A., 129 Galáz, J. See Sielfeld, W., 336
Fritsch, F.E., 79 Galáz, L.E. See Camus, P.A., 309
Fromentin, J.M. See Walther, G.R., 476 Galbraith, H., 461
Froneman, P.W., 129 Galeron, J. See Fabri, M.C., 167
See Teske, P.R., 339 Gales, R. See Bunce, A., 456
Frost, B.W. See Bollens, S.M., 455 See Pemberton, D., 470
See Hays, G.C., 130 Gall, A. See Mumby, P.J., 469
Frost, M. See Martin, D., 401 Gall, M. See Boyd, P.W., 456
See Moschella, P.S., 402 Gallardo, V. See Milessi, A., 327
495
AUTHOR INDEX
Gallardo, V.A., 316 Gaylord, B., 79, 461

See Arntz, W., 306 See Denny, M., 458
See Atkinson, L.P., 306 See Raimondi, P.T., 84
See Carrasco, F.D., 309 See Reed, D.C., 85
See Escribano, R., 314 See Siegel, D.A., 336
See Fossing, H., 316 Gaymer, C.F. See Brokordt, K.B., 308
See Gutiérrez, D., 319 See Thiel, M., 199–344
See Neira, C., 329 Geesey, M.E. See Hutchins, D.A., 321
See Palma, M., 331 Gelcich, S., 317
See Roa, R., 333 See Castilla, J.C., 310
See Schulz, H.N., 335 Gelfman, C. See Wishner, K.F., 344
See Sellanes, J., 336 Geller, P.E., 34
Gallegos, M.E. See Marbà, N., 401 Gellner, G. See Rooney, N., 135
Galleguillos, R., 316 Genin, A., 129, 461
See Chong, J., 311 See Haury, L., 130
See Gómez-Uchida, D., 317 Genner, M.J. See Sims, D.W., 473
Galois, R. See David, V., 127 Gentili, B. See Morel, A., 35
Gama, A.M. da S., 129 Gentleman, W. See Miller, C.B., 133
Ganade, G. See Fonseca, C.R., 460 George, R. See Guinotte, J.M., 462
Gandini, P. See Frere, E., 316 Georges, J.Y. See Ferraroli, S., 460
Gao, K., 461 Geraci, S. See Fanelli, G., 398
Gao, K.S., 461 Gerard, V.A., 79
Garcés-Vargas, J. See Fuenzalida, R., 316 See Bruhn, J., 456
García, C. See Buschmann, A.H., 75, 308 See Burgman, M.A., 75
Garcia, C.M., 461 Gerber, L.R. See Micheli, F. 327
Garcia, H.E. See Conkright, M.E., 311 Gerdes, D. See Palma, M., 331
García, J.C., 317 Gering, J.C. See Veech, J.A., 172
Garcia, O. See Hernández-Carmona, G., 80 Gerlotto, F. See Bertrand, A., 307
Garcia-de la Rosa, O. See Hernández-Carmona, G., 81 Gerrodette, T. See Dayton, P.K., 76, 313
García-González, D. See Drake, P., 128 Gershwin, L.A., 461
Garde, K., 461 Ghelardi, R.J., 79
See Wängberg, S.A., 476 Giannoulaki, M. See Karakassis, I., 169
Gardner, N.L. See Setchell, W.A., 86 Gibb, J.G. See de Lange, W.P., 458
Gardner, T.A., 461 Gibb, S.W. See Brown, B.E., 191
Garkaklis, M.J. See Bancroft, W.J., 454 Gibbons, J. See Capella, J., 309
Garman, G.D., 79 Giberto, D.A. See Schiariti, A., 135
Garnett, S.T., 461 Gibson, G. See Hu, Y.-X., 34
Garrabou, J., 461 Gibson, R.N., 129
See Perez, T., 470 Giesecke, R., 317
Garrido, J. See González, J., 318 See Escribano, R., 314
Garrison, L.P. See Link, J.S., 132 See González, H.E., 317
Garthe, S. See Ludynia, K., 324 Gieskes, W.W.C. See Kelfkens, G., 465
See Simeone, A., 337 Giglio, S. See Marchant, M., 325
See Weichler, T., 343 Gilbert, J.J. See Twining, B.S., 137
Garwood, P.R. See Frid, C.L., 316 Gili, J.M. See Pagès, F., 330
Gaspar, P. See Ferraroli, S., 460 See Ribes, M., 134
Gasparovic, F. See Jeftic, L., 400 Gill, J.A. See Fish, M.R., 460
Gassmann, G. See Maier, I., 83 See Gardner, T.A., 461
Gaston, K.J., 168 See Smart, J., 473
See Spicer, J.L., 337 Gillanders, B.M. See Beck, M.W., 394
See Williams, P.H., 172 See Bloomfield, A.L., 455
See Williamson, M., See Fowler-Walker, M.J., 460
Gates, R.D. See Gleason, D.F., 461 Gillett, N.P., 461
Gatien, G. See Freeland, H.J., 316 Gilman, S.E. See Barry, J.P., 454
Gatlin, D.M., III, 129 See Sagarin, R.D., 472
Gattuso, J.P., 461 Gilmour, J., 461
See Engel, A., 460 Giordano, M., 461
See Kleypas, J.A., 466 Giraldo, A., 317
Gaughan, D.J., 461 Girondot, M. See Rivalan, P., 472
496
AUTHOR INDEX
Gittleman, J.L. See Mace, G.M., 169 See Giesecke, R., 317
Glascock, M.D. See Sandweiss, D.H., 334 See Grünewald A., 318
Glatts, R.C. See Kaufmann, R.S., 131 See Iriarte, J.L., 321
Gleason, D.F., 461 See Marchant, M., 325
Glemaréc, M., 398 See Morales, C.E., 328
See Barbera, C., 394 See Pagès, F., 330
Glen, F., 461 See Pantoja, S., 331
See Hays, G.C., 463 See Pavez, M.A., 331
Gloersen, K.A. See White, W.B., 476 See Rodríguez-Graña, L., 333
Glover, A.G., 461 See Thiel, M., 199–344
Glover, H.E. See Codispoti, L.A., 311 See Vargas, C.A., 341
Glud, R.N., 317 González, J., 318
See Fossing, H., 316 González, L.A., 129
Glue, D.F. See Crick, H.Q.P., 457 González, M. See Jaramillo, E., 321
Glynn, P.W., 317 González, R.R., 318
Gnanadesikan, A. See Griffies, S.M., 462 See Ulloa, O., 340
See Orr, J.C., 470 See Yaikin, J., 344
Gobin, J.F., 168 González, S.A., 318
Goddard, J. See Langdon, C., 466 See Stotz, W., 338
Goddard, S.M., 129 Gonzalez-Fragoso, J., 79
Godfrey, J.S. See Ridgway, K.R., 472 Gonzalez-Rodas, G. See El-Sayed, S., 459
See Tomczak, M., 194 Goodall, J. See Dunton, K., 313
Godfrey, M.H., 461462 Gooday, A.J., 318
Godínez-Orta, L. See Takesue, R.K., 339 See Levin, L.A., 169, 324
Godley, B.J. See Broderick, A.C., 456 Goodson, M.S., 191
See Hays, G.C., 463 Gorczynska, M.I. See Mumby, P.J., 469
Godo, O.R. See Byrkjedal, I., 456 Gordillo, S. See Adami, M.L., 74
Godoy, N.E., 317 Gordon, H.B., 462
Goericke, R. See Moore, L.R., 469 Gordon, H.R., 34
Goes, J.I., 462 Gordon, J. See Davidson, N.C., 396
Goff, L.J. See Garman, G.D., 79 Gordon, R.M. See Martin, J.H., 326
Goffart A, 462 Gorelick, R. 168
Goggin, C.L., 462 Gorgula, S.K., 462
Goldberg, N.A., 462 Gorshkov, S.G., 318
Golding, N. See Connor, D.W., 396 Gotelli, N.J., 168
Goldsmith, T.H. See Losey, G.S., 467 See Colwell, R.K., 167
Gomez, E.D. See Kaufman L., 322 Gowing, M.M. See Wishner, K.F., 344
Gómez, F. See Bertrand, A., 307 Gowlett-Holmes, K. See Koslow, J.A., 466
Gómez, I., 317 See Williams, A., 476
See Huovinen, P., 321 Graco, M., 318
See Thiel, M., 199–344 See Farías, L., 315
See Véliz, K., 342 Graddon, D.J. See Barrett, N.S., 397
See Westermeier, R., 343 Graf, G., 318
Gomez, M. See Calliari, D., 126 Graham, M.H., 39–88, 79, 80, 318, 462
Gómez-Uchida, D., 317 See Erlandson, J.M., 78
Gomoiu, M.-T., 398 See Hernández-Carmona, G., 80
Gong, D.Y., 317 See Kinlan, B.P., 82
Gong, W.K. See Farnsworth, E.J., 460 See Munoz, V., 83, 329
González, A., 317 See Muñoz, V.,
See Leibold, M.A., 323 See Sala, E., 85
González, A.E. See Thiel, M., 199–344 See Steneck, R.S., 86, 404, 474
González, C., 317 Graham, N.E. See Polovina, J.J., 471
See Fernández, M., 315 Graham, W.M. See Reed, D.C., 85
See Ibáñez, C.M., 321 Grall, J., 398
González, E. See Thiel, M., 339 See Barbera, C., 394
González, E.O. See Camus, P.A., 309 See Hall-Spencer, J.M., 398
González, H. See Bernal, P.A., 307 Grams, G.W. See Chýlek, P., 34
González, H.E., 317, 318 Granados, I. See Takesue, R.K., 339
See Escribano, R., 314 Graneli, E. See Carlsson, P., 456
497
AUTHOR INDEX
Granhag, L. See Jonsson, P.R., 465 Guerrero, R.A. See Schiariti, A., 135
See Moschella, P.S., 402 Guidetti, P., 398
Grant, A. See Gardner, T.A., 461 See Rismondo, A., 403
Grantham, B.A., 318 Guiler, E.R., 80
See Shanks, A.L., 336 Guillén, O. See Zuta, S., 344
Grassl, H. See Pozdnyakov, D., 35 Guinéz, R. See Buschmann, A.H., 308
Grassle, J.F., 168 See Castilla, J.C., 310
Grasso, M. See Costanza, R., 396, 457 See Alvarado, J.L., 305
Grau, A. See Delgado, O., 396 See Kaplan, D.M., 322
Grau, A.M. See Meinesz, A., 401 Guinotte, J. See Kleypas, J.A., 466
Graves, J.E., 318 Guinotte, J.M., 462
Gravestock, P. See Balmford, A., 307 Guisado, C. See Véliz, D., 342
Gray, J.S. 168, 398 Guizien, K., 318
See Ellingsen, K.E., 167 Gullström, L.B. See Baden, S., 394
See Ugland, K. I., 172 Gundersen, J.K. See Fossing, H., 316
Grebmeier, J. See Dunton, K., 313 See Glud, R.N., 317
Green, E.P., 398 Gunn, J. See Polacheck, T., 471
See Spalding, M.D., 194, 404 Gunn, J.S. See Thresher, R.E., 474
Green, K., 129 Gunther, E.R., 318
Green, M.J.B., 398 Günther, C.-P. See Grimm, V., 129
Green, R.E. See Balmford, A., 394 Günther, R.T., 398
Greene, C.H. See Gal, G., 129 Guppy, H.B., 319
See Smith, S.L., 136 Gupta, T.D. See Pal, T., 192
Greenlaw, C.F., 129 Gurevitch, J., 168
Gregory, J.M., 462 See Rosenberg, M.S., 171
See Church J.A., 457 Gurney, W. See Speirs, D., 136
Greig, A. See Beare, D.J., 454 Gurney, W.S.C. See Lawrie, S.M., 132
Grelle, V. See Eduardo, C., 167 Gustafsson, L. See Both, C., 455
Greve, W., 462 Gustavson, K. See Wängberg, S.A., 476
See Bonnet, D., 455 Gutierrez, A., 80, 319
See Winkler, G., 138 Gutiérrez, D. See Arntz W.E., 306
Griffies, S.M., 462 See Escribano, R., 314
Griffin, D.A., 462 See Gallardo, V.A., 316
Griffiths, F.B. See Harris, G.P., 463 See Graco, M., 318
See Sedwick, P.N., 473 See Muñoz, P., 329
Griffiths, R.C. See Rao, T.S.S., 193 See Neira, C., 329
Griffiths, S.P., 462 See Sellanes, J., 336
Grimes, D.J. See Harvell, C.D., 463 See Tarazona , J., 339
Grimm, V., 129 Gutow, L. See Franke, H.D., 398
See Peña T.S., 331 See Thiel, M., 87, 339
Grol, M.G.G. See Dorenbosch, M., 459 Guzman, H.M. See Pandolfi, J.M., 470
Grone, G. See Rodríguez, L., 333 Guzmán, N. See Ortlieb, L., 330
Grootes, P.M. See Haneburth, T., 192 See Takesue, R.K., 339
Grosberg, R. See Hughes, T.P., 464 Guzman-Speziale, M., 191
Grove, R.S., 462 Gwak, W.-S. See Tanaka, Y., 136
Grover, J. 462
Grua, P., 80
Gruber, A. See Wiencke, C., 476 H
Gruber, N. See Orr, J.C., 470
Grünewald A., 318 Haaker, P.L. See Davis, G., 458
Grzybowski, K. See Simeone, A., 337 Häder, D., 462
Gucinski, H. See Behrenfeld, M., 455 Häder, D.P., 462
Guderley, H. See Martínez, G., 326 Häder, M.A. See Häder, D., 462
Guderley, H.E. See Brokordt, K.B., 308 Hadfield, M. See Boyd, P.W., 456
Guerao, G. See Delgado, L., 127 Hagen, N.T., 398
Guerin-Ancey, O. See David, P.M., 127 Haight, W.R. See Polovina, J.J., 471
Guerra, C., 318 Halim, Y. See Jeftic, L., 400
See Sielfeld, W., 336 Hall, J. See Boyd, P.W., 456
Guerra, Y. See Sielfeld, W., 336 Hall, J.A. See Frid, C.L.J., 398
498
AUTHOR INDEX
Hall, K.R. See Root, T.L., 472 Harmelin-Vivien, M.L. See Salen-Picard, C., 334
Hall, R., 192 Harper, J.L., 168
Hall, S.J., 462 Harrington, B. See Galbraith, H., 461
See Frid, C.L.J., 398 Harris, G., 463
Hallacher, L.E., 80 See Boersma, D., 307
Hällfors, G. See Kangas, P., 400 Harris, G.P., 463
Halling, C. See Chopin, T., 311 See Thresher, R.E., 474
Hall-Spencer, J., 398 Harris, G.W. See Procter, T.D., 35
Hall-Spencer, J.M., 399 Harris, L.G., 80
See Barbera, C., 394 Harris, R. See Bonnet, D., 455
See Grall, J., 398 Harrison, P.J. See Hurd, C.L., 81
Halpern, B. See Beck, M.W., 394 Harrison, P.L. See Babcock, R.C., 454
Halpern, B.S., 80 Harrison, W.G. See Longhurst, A.R., 132
See Graham, M.H., 79 Harrold, C., 80, 319
See Roberts, C.M., 333 See Graham, M.H., 79
Halpin, P.M., 319 See Watanabe, J.M., 88
Hama, T. See Goes, J.I., 462 Hartmann, G. See Hartmann-Schröder, G., 319
Hamann, J. See Wilhelm, F.M., 138 Hartmann, K. See Hobday, A.J., 464
Hamerlynck, O. See Mees, J., 133 Hartmann-Schröder, G., 319
Hamilton, A.J., 168 Hartnoll, R.G. See Oh, C.W., 133
Hamm, L., 399 Harty, C., 463
Hammer, C. See Frid, C., 398 Harvell, C.D., 463
Hamner, W.M. See Carlton, J.H., 126 Harvey, M. See Boyd, P.W., 456
See Omori, M., 134 Harwood, K.G. See Frid, C.L.J., 398
Hampel, H., 129 Harzhauser, M., 319
Hampson, G.R. See Sanders, H.L., 171 Hastings, A. See Botsford, L.W., 307
Hampton, J. See Lehodey, P., 467 See Lockwood, D.R., 324
Hamre, B. See Erga, S.R., 460 See McCann, K., 133
Hamren, U., 130 Hatchwell, B.J. See Votier, S.C., 475
Hanamura, Y., 130 Hatton, C. See Thrush, S.F., 474
Handa, N. See Goes, J.I., 462 Haury, L., 130, 319
Haneburth, T., 192 Hauser, L. See Gómez-Uchida, D., 317
Hanelt, D., 462 Hauxwell, J., 399
See Bischof, K., 455 Hawkes, M.W. See Gabrielson, P.W., 79
Hannach, G., 319 Hawkins, S. See Hiscock, K., 464
Hannah, L., 399 Hawkins, S.J.,
Hannemann, A. See Behrenfeld, M., 455 See Airoldi, L., 394
Hannon, B. See Costanza, R., 396, 457 See Jonsson, P.R., 465
Hansen, J., 462 See Kendall, M.A., 465
Hansen, L.J. See Roessig, J.M., 472 See Martin, D., 401
Hansen, O.S. See Nielsen, T.G., 192 See Mieszkowska, N., 469
Hansen, P.J. See Nielsen, T.G., 192 See Moschella, P.S., 402
See Pedersen, M.F., 470 See Sims, D.W., 473
Hanson, J.M., 130 See Southward, A.J., 474
Hansson, S., 130 See Thompson, R.C., 404
See Hamren, U., 130 Hawskworth, D.L. See Harper, J.L., 168
See Rudstam, L.G., 135 Haxo, F.T. See Neushul, M., 83
Harborne, A.R., 463 Hay, C.H., 80
See Mumby, P.J., 469 Haye, P. See Thiel, M., 339
Harch, B.D. See Manson, F.J., 468 Haye, P.A. See Thiel, M., 199–344
Hardy, J. See Behrenfeld, M., 455 Hayek, L.C., 168
Hare, C.E. See Hutchins, D.A., 321 See Buzas, M.A., 166
Hare, S.R. See Mantua, N.J., 468 Hays, C. See Duffy, D., 313
Harger, B.W.W. See Lewis, R., 82 Hays, C.G. See Beck, M.W., 394
See Nyman, M.A., 84 Hays, G.C., 130, 463
Hargraves, P. See Keller, A.A., 465 See Broderick, A.C., 456
Harlay, J. See Engel, A., 460 See Luschi, P., 467
Harmelin, J.G. See Garrabou, J., 461 See McMahon, C.R., 468
See Perez, T., 470 Hayward, T., 463
499
AUTHOR INDEX
Hayward, T.L. See Venrick, E.L., 475 Herman, J.S. See MacLeod, C.D., 468
Haywood, M.D.E. See Vance, D.J., 475 Herman, P.M.J. See Dauwe, B., 127
Head, E.J.H. See Longhurst, A.R., 132 See Thrush, S.F., 474
Heads, M.J. See Chin, N.K.M., 76 Herman, S.S., 130
Heales, D.S. See Vance, D.J., 475 Hermosilla, C. See Zagal, C., 344
Healy, M.G., 399 Hermosilla, N. See Jerardino, A., 322
Heard, R. See Thiel, M., 339 Hernández, C. See Bertrand, A., 307
Heard, R.W. See Price, W.W., 134 Hernández, C.E., 319
Hebbeln, D., .319 See Cárdenas, L., 309
See Antia, A.N., 454 See Moreno, R.A., 328
See Coloma, C., 311 See Poulin, E., 332
See González, H.E., 317 Hernández, E. See Narváez, D.A., 329
See Marchant, M., 325 See Poulin, E., 332
See Muñoz, P., 329 Hernández, E.H., 319
See Palma, M., 331 See Castro, L.R., 311
Heck, K.L., Jr, 399 Hernández-Carmona, G., 80, 81
See Beck, M.W., 394 See Edwards, M.S., 78
See Mattila, J., 132 See Ladah, L.B., 82, 323
Hecq, J.H. See Goffart, A., 462 Hernández-González, M.C. See Buschmann, A.H., 75, 308
Hector, A. See Purvis, A., 170 See Muñoz, V., 83, 329
Hedgepeth, J.W., 168 Hernández-Miranda, E., 319
Hedges, L.V., 168 Hernando, M.P., 463
See Gurevitch, J., 168 Herrera, G. See Llanos, A., 324
Hedley, R.H. See Buchanan, J.B., 126 See Llanos-Rivera, A., 324
Heemann, C. See Engel, A., 460 Herrera, G.A. See Landaeta, M.F., 323
Heffels, C.M.G., 34 Herrera, L., 319
Heidemann, A. See Valdivia, N., 340 Herring, S. See Boss, E., 33
Heine, J.N. See Spalding, H., 86 Herring, S.G., 34
See Spalding, H., 474 Herrod-Julius, S. See Galbraith, H., 461
Heinemeier, J. See Petersen, K.S., 403 Hertweck, C. See Maier, I., 83
Heino, M. See Byrkjedal, I., 456 Herzka, S.Z., 319
Heinrich, A.K., 319 Hesp, P.A., 192
Heinrich, D. See Lotze, H.K., 401 Hess, S. See Gómez, I., 317
Heip, C.H.R. See Dauwe, B., 127 Hessen, D., 463
Heitzmann, D. See Heffels, C.M.G., 34 Hesslein, R.H. See Johannsson, O.E., 130
Helbling, W. See Barbieri, E.S., 454 Hessler, R.R. See Levin, L.A., 169, 324
HELCOM, 399 See Sanders, H.L., 171
Hell, R. See Giordano, M., 461 Hewavisenthi, S., 463
Hellberg, M.E. See Baums, I.B., 307 Hewitt, J.E. See Lohrer, A.M., 467
See Swearer, S.E., 339 See Norkko, A., 470
Helly, J.J., 319 See Thrush, S.F., 171, 474, 475
Helmuth, B., 463 See Turner, S.J., 404
Helmuth, B.S., 80 Heyen, H. See Kroencke, I., 323
Hemminga, M.A., 399 Heyward, A. See Negri, A., 469
Hempel, W.M., 80 Heyward, A.J. See Babcock, R.C., 454
See Eardley, D.D., 78 Hickey, B.M. See Hill, A.E., 320
Henderson, P.A. See Bamber, R.N., 125 Hickey, K.R. See Healy, M.G., 399
Hendon, H. See Allan, R., 190 Hickling, R., 463
Hennicke, J.C., 319 Hicks, J.T. See King, B.R., 466
Hennigar, A.W. See Scheibling, R.E., 473 Hidalgo, P., 320
Henríquez, L.A. See Buschmann, A.H., 308 See Escribano, R., 314
Henry, E.C., 80 See Fernández, D., 315
Henry, G.W., 463 See Marín, V.H., 326
See Lyle, J.M., 468 Higgins, K. See Botsford, L.W., 307
Herbert, R.J. See Mieszkowska, N., 469 Hilborn, R., 320
Herbold, B. See Feyrer, F., 128 Hildebrandt, H. See Grimm, V., 129
Herbst, L., 463 Hill, A.E., 320
Herbst, L.H. See Ene, A., 460 Hill, J.K. See Hickling, R., 463
Herdiana, Y. See Baird, A.H., 190 See Parmesan, C., 470
500
AUTHOR INDEX
Hill, M., 168 Hogue, V.E., 464

Hill, M.O. See Davies, C.E., 396 Holben, B.N. See Dubovik, O., 34
Hillebrand, H., 34, 168 See Zhao, T.X.-P., 36
Hilse, C. See Wilhelm, C., 476 Holberton, R. See Helmuth, B.S., 80
Hilton, M. See Hesp, P.A., 192 Holbrook, S.J., 81, 464
Hilton-Taylors, C. See Brooks, T.M., 395 Holby, O. See Glud, R.N., 317
Himmelman, J.H. See Brokordt, K.B., 308 Holland, G.J. See Webster, P.J., 476
Hines, A.H. See Pearse, J.S., 84 Hollenberg, G.J. See Abbott, I.A., 74
Hinga, K.R., 464 Holley, M.C. See Brown, B.E., 191
Hinkley, S. See MacNally, R., 169 Holliday, D.V. See Greenlaw, C.F., 129
Hinojosa, I., 320 See Kringel, K., 131
Hinojosa, I.A. See Macaya, E.C., 83, 325 Holm, P. See Lotze, H.K., 401
Hinojosa, L.F., 320 Holme, N.A., 168
Hirose, T. See Takahashi, K., 136 See Eleftheriou, A., 128
Hirota, Y., 130 Holmer, M., 399
Hirst, A. See Bonnet, D., 455 See Marbà, N., 401
Hirst, A.C. See Gordon, H.B., 462 Holmes, K.E. See Harborne, A.R., 463
See Sarmiento, J.L., 473 Holm-Hansen, O. See Mitchell, B.G., 469
Hirst, A.G., 464 Holst, M. See Boulding, E.G., 455
Hiscock, K., 399, 464 Holt, G.J. See Herzka, S.Z., 319
See Mieszkowska, N., 469 Holt, R.D. See Leibold, M.A., 323
Hitchcock, D.R. See Newell, R.C., 402 Holt, S.A. See Herzka, S.Z., 319
Hobart, W.L. See Mackenzie, C.L., Jr, 401 Holthuis, L.B. See van der Baan, S.M., 137
Hobbie, J.E. See Hansson, S., 130 Holyoak, M., 130
Hobday, A. See Campbell, R.A., 456 See Leibold, M.A., 323
See Polacheck, T., 471 Hong, S.Y., 130
Hobday, A.J., 81 Hooff, R. See Shanks, A.L., 86
See Koslow, J.A., 466 Hooke, J.M. See Bray, M.J., 395
See Poloczanska, E.S., 407–478, 464 Hooker, J.D., 81
Hobson, E.S., 81 Hooker, Y. See Lleellish, J., 324
Hock, L. See Alpers, W., 190 See Lleellish, M., 83
Hocking, P.J. See Cambridge, M.L., 456 Hoopes, M.F. See Leibold, M.A., 323
Hockley, N. See Balmford, A., 307 Hoovenier, J.W., 34
Hodkinson, J. R., 34 See Mishchenko, M.I., 35
Hoedt, F. See Ward, T.M., 476 See Volten, H., 36
Hoegh-Guldberg, H., 464 Hoozemans, F.M.J. See Nicholls, R.J., 402
Hoegh-Guldberg, O., 192 Hope, A.S. See Augenstein, E.W., 74
See Donner, S.D., 459 Hopkins, T.L., 130
See Hoegh-Guldberg, H., 464 Hore, J.P., 399
See Hughes, T.P., 464 Horgan, E.F. See Brown, H., 126
See Jones, R.J., 465 Hormazábal, S., 320
See Kleypas, J.A., 466 See Morales, C.E., 328
See LaJeunesse, T.C., 192 See Rutllant, J.A., 334
See Poloczanska, E.S., 407–478, 464 See Santelices, B., 335
See Raven, J., 471 See Shaffer, G., 336
See Walther, G.R., 476 See Ulloa, O., 340
Hoerling, M.P. See Diaz, H.F., 77 See Yuras, G., 344
Hoffmann, A., 320 Horvat, M. See Trombini, C., 404
See Santelices, B., 334 Horvath, H. See Kocifaj, M., 35
Hoffmann, A.J., 81, 320 Hoschino, K. See Beck, M.W., 394
See Avila, M., 75 Hostens, K., 130
See Santelices, B., 85 Hostetler, C. See Hu, Y.-X., 34
Hoffmann, L. See Engel, A., 460 Houghton, J.D. See Hays, G.C., 463
Hoffmann, R.C., 399 Houk, J.L. See McCleneghan, K., 83
See Lotze, H.K., 401 Howe, M.A., 81
Hoffmann, S. See Greve, W., 462 Howell, E.A. See Polovina, J.J., 471
Hoffmeyer, M.S., 130 Howell, K.L. See Connor, D.W., 396
Hofmann, E.E. See Harvell, C.D., 463 Howlett, B.G. See Boulter, S.L., 455
Hofmann, G.E., 320 Hu, Y.-X., 34
501
AUTHOR INDEX
Huang, C.Y. See Lee, K.K., 467 Hüttel, M. See Fossing, H., 316
Huang, M., 130 Huybrechts, P. See Church J.A., 457
Hubbard, D.M. See Dugan, J.E., 313 Huyer, A., 321
Hubbell, S.P., 168 See Freeland, H.J., 316
Hubbs, C.L. See North, W.J., 84 See Grantham, B.A., 318
Hucke-Gaete, R. See Moreno, C.A., 328 See Strub, P.T., 338
Hückstädt, L.A., 320 Hyde, D. See Losey, G.S., 467
Hudson, G. See Menge, B.A., 327 Hyder, P., 192
Hudson, I.L. See Keatley, M.R., 465 Hylleberg, J. See Janekarn, V., 192
Huesemann, M.H., 464
Huettel, M. See Wild, C., 343
Huffman, D.R. See Bohren, C.F., 33 I
Huges, T.P. See Jackson, J.B.C., 399
Ianora, A., 321
Huggett, C.L. See Levin, L.A., 324
Ibanez, F. See Beaugrand, G., 454
Huggett, J.A. See Verheye, H.M., 342
Ibáñez, C. See Chong, J., 311
Hughes, A.R. See Byrnes, J., 75
Ibáñez, C.M., 321
Hughes, B. See Hernández-Carmona, G., 80
Ibáñez, S. See Moreno, C.A., 328
Hughes, D.J. See Davison, D.M., 396
Ibarra-Obando, S.E. See Gonzalez-Fragoso, J., 79
Hughes, J.M.R. See Jones, T.A., 400
ICES, 399
Hughes, L., 464
IFOP, 321
See Beaumont, L.J., 454
Iglesias-Prieto, R. See Kleypas, J.A., 466
See Chambers, L.E., 457
Illanes, J.E., 321
Hughes, R.G., 399
Hughes, T.P., 464
Ilyasd, M. See McKenzie, R.L., 468
See Pandolfi, J.M., 402, 470
IMARPE, 321
Huisman, J., 464
Immonen, A.K. See Markkula, S.E., 468
Hulbert, E.M., 130
Inagaki, H. See Ohtsuka, S., 134
Hull, P.G. See Quinby-Hunt, M.S., 36
Indacochea, A. See Tarazona, J., 339
Hultgren, K.M. See Byrnes, J., 75
Inglis, G., 81
Humboldt, A., 320
Inglis, G.J., 464
Humphrey, C. See Negri, A., 469
Inoue, T., 130
Hung, C.C. See Hesp, P.A., 192
Invers, O., 464
Hung, O.S. See Chan, B.K.K., 457
Iotti, M. See See Simonini, R., 404
Hunt, A.J. See Quinby-Hunt, M.S., 36
IPCC, 192, 464
Hunt, G.L. See Jahncke, J., 321
Iratchet, P. See Ortlieb, L., 330
Hunt, J.W. See Anderson, B.S., 74
Iriarte, J. See González, H.E., 317
Hunter, C.N. See Martin, J.H., 326
Iriarte, J.L., 321
Hunter, F.M. See Votier, S.C., 475
See González, H.E., 318
Hunter, J.R., 464
See Pizarro, G., 332
See Porteiro C., 471
Hunter, K.A. See Jickells, T.D., 465
Irribarren, C. See Escribano, R., 314
Huntley, B. See Parmesan, C., 470
See Rodríguez, L., 333
Huntley, M., 464
Isaac, W.E., 81
Huovinen, P., 321
Ishida, A. See Orr, J.C., 470
See Gómez, I., 317
Ishihara, T. See Gao, K., 461
Huovinen, P.J., 81
Ishimatsu, A. See Kikkawa, T., 466
Hurd, C.L., 81
Isla, E., 306
Hurd, S.D. See Polis, G.A., 332
Islam, M.S., 399
Hurlbert, A.H., 168
Ittekkot, V. See Burkett, V., 456
Hurlbert, S., 168
IUCN, 321
Hurlbert, S.N., 130
Ivankina, E.V. See Both, C., 455
Hurrel, J.W. See Trenberth, K.E., 340
Ivanov, V.V. See Beamish, R.J., 454
Hussain, S.A. See Badola, R., 454
Ivesa, L. See Meinesz, A., 401
Huston, M. A., 168
Hutchings, L. See Verheye, H.M., 342
Hutchins, D.A., 321 J
See Sedwick, P.N., 473
Hutchinson, C. See Hannah, L., 399 Jablonski, D., 169
Hutson, A.M. See Robinson, R.A., 472 See Roy, K., 171
502
AUTHOR INDEX
Jackett, D.R. See Gregory, J.M., 462 Jaubert, J.M., 400

See Walsh, K.J.E., 476 Jeans, D.R.G. See Hyder, P., 192
Jackson, A., 399 Jefferies, R.L. See Wolters, M., 405
Jackson, C.J., 130 Jeftic, L., 400
Jackson, C.T. See Polis, G.A., 332 Jelinski, D.E. See Mulkins, L.M., 133
Jackson, D. See Báez, P., 306 See Orr, M., 84
Jackson, D.G. See Méndez, C.A., 327 Jeltsch, F. See Grimm, V., 129
Jackson, G. See Ward, T.M., 476 Jenkins, G.P. 464
Jackson, G.A., 81 Jenkins, M. See Balmford, A., 394
See North, W.J., 84 Jeno, K. See Fernández, M., 315
Jackson, J.B. See Kaufman L., 322 Jensen, A., 400
See Pandolfi, J.M., 470 Jensen, C. See Lotze, H.K., 401
Jackson, J.B.C., 399 Jensen, J.N. See Rumohr, H., 171
See Hughes, T.P., 464 Jensen, J.R., 81
See Lotze, H.K., 401 Jensen, M.H. See Rask, N., 403
See Pandolfi, J.M., 402 Jeon, H. See Keller, A.A., 465
Jacob, B. See Cuevas, L.A., 312 Jeppesen, E. See Aaser, H., 124
See Daneri, G., 312 Jerardino, A., 322
See Troncoso, V.A., 340 Jerez, G. See Aviléz, O., 306
Jacobs, J.R. See Potts, D.C., 193 Jerling, H.L., 130
Jacobsen, F.R. See Dean, T.A., 77 Jerlov, N.G., 34, 464
Jacobson L.D. See Porteiro C., 471 Jernakoff, P., 465
Jacquemin, B. See Rivalan, P., 472 Jesse, S., 322
Jacquet, S. See Engel, A., 460
See Ortiz, M., 330
Jaenicke, L. See Maier, I., 83
Jesus, L. See Azeiteiro, U.M.M., 125
Jaffe, J.S. See De Robertis, A., 128
Jetten, M.S.M. See Kuypers, M.M., 323
Jagannathan, S. See Makris, N.C., 132
Jeudy de Grissac, A. See Boudouresque, C.F., 395
Jahncke, J., 321
Jex, E. See Hore, J.P., 399
Jaklin, A. See Meinesz, A., 401
Ji, Y. See Gao, K.S., 461
See Zavodnik, N., 405
Jickells, T.D., 465
Jaksic, F.M. See Castro, S.A., 311
Jiménez, J.E. See Buschmann, A.H., 308
Jakubauskas, M.E. See Su, J.C., 171
Jiménez, M., 322
James, C. See Strub, P.T., 338
Jo, S.-G. See Hanamura, Y., 130
James, D.E. North, W.J., 84
See Suh, H.-L., 136
Jameson, G. See Boyd, P.W., 456
Jodwalis, C.M. See Dunton, K.H., 78
Jameson, R.J. See Laidre, K.L., 82
Janekarn, V., 192 Johannsson, O.E., 130
See Nielsen, T.G., 192 Johanssen, A. See Brattström, H., 308
Janodet, E. See Blasco, F., 455 Johansson, G., 400
Janssen, J., 130 See Eriksson, B.K., 398
Jansson, B.-O., 400 Johansson, J. See Wahl, M., 475
Jantarapagdee, P. See Yesaki, M., 194 Johansson, S. See Hansson, S., 130
Jara, A.F. See Moreno, C.A., 328 See Rudstam, L.G., 135
Jara, C.G., 321 Johns, D.G. See Edwards, M.J., 459
Jara, F., 321 Johnson, C., 465
Jara, H.F., 321 Johnson, C.R. See Valentine, J.P., 475
See Moreno C.A., 83 Johnson, D.D., 322
Jaramillo, E., 321, 322 Johnson, D.R., 322
See Contreras, H., 312 Johnson, E.J. See Leet W.S., 82
See Duarte, C., 313 Johnson, K.A., 400
See Dugan, J.E., 313 Johnson, K.S. See Coale, K.H., 311
See Fernández, M., 315 Johnson, L.E. See Brawley, S.H., 456
See McLachlan, A., 326 Johnson, N.A. See Rex, M.A., 170
See Quijón, P., 332 Johnson, T. See Peterson, W.T., 331
See Stotz, W., 338 Johnson, W.S., 130
Jarre, A. See Cury, P., 312 Johnston, D.W. See Norse, E.A., 170
See Moloney, C.L., 327 Johnston, N.M., 130
Jarvinen, A. See Both, C., 455 Johnstone, I., 192
Jatczak, K. See Menzel, A., 469 Johst, K. See Peña T.S., 331
503
AUTHOR INDEX
Jokiel, P.L. See Kleypas, J.A., 466 Kadyshevich, Ye. A., 34

See Shick, J.M., 473 Kaehler, S., 82
Jokinen, E.M. See Markkula, S.E., 468 Kaila, L. See Parmesan, C., 470
Jonasz, M., 34 Kain, J., 322
Jones, A.G. 465 Kain, J.M., 82
Jones, A.P. See Fish, M.R., 460 Kaiser, B. See Wilhelm, C., 476
Jones, A.R. See Kokhanovsky, A.A., 35 Kaiser, M.J. See Gelcich, S., 317
Jones, C.G., 322 Kamezaki, N. See Bowen, B.W., 455
Jones, E.G. See Beare, D.J., 454 Kamykowski, D., 322
Jones, G.P. See Swearer, S.E., 339 Kangas, P., 400
Jones, J.M., 465 Kaplan, D. See Velázquez, I., 342
Jones, K. See Ward, T.M., 476 Kaplan, D.M., 322
Jones, L.G., 82 See Wieters, E.A., 343
See North, W.J., 84 Kappel, C.V. See Pandolfi, J.M., 470
Jones, M.B. See Mees, J., 133 Karagatzides, J.D. See Mulkins, L.M., 133
See Moffat, A.M., 133 Karakassis, I., 169
See Roast, S.D., 135 See Rumohr, H., 171
Jones, N.S. 169 Karanas, J.J., 465
See Gage, J., 167 Karlson, K., 400
Karlsson, L. See Jonzén, N., 465
Jones, R. See Done, T.J., 459
Karoly, D.J., 465
See Galbraith, H., 461
Karp-Boss, L., 35
See McEvedy, C., 401
See Clavano, W.R., 1–38
See Negri, A., 469
Kartawijaya,T. See Baird, A.H., 190
Jones, R.J., 465
Kaska, D. See Hempel, W.M., 80
Jones, T.A., 400
Kassem, K. See Burke, L., 396
Jones, W.R. See Janssen, J., 130
Kastendiek, J. See Dixon, J., 77
Jonsson, B.G. See Sih, A., 404
Kathiresan, K., 465
Jonsson, P. See Rosenberg, R., 403
See Moorthy, P., 469
Jonsson, P.R., 465 Kaufman L., 322
See Airoldi, L., 394 Kaufmann, R.S., 131
See Moschella, P.S., 402 Kaupp, S.E. See Hunter, J.R., 464
Jonzén, N., 465 Kausch, H. See Köpcke, B., 131
Joos, F. See Orr, J.C., 470 Kautsky, H. See Kautsky, N., 400
Jordi, A. See Marbà, N., 401 Kautsky, L. See Serrão, E.A., 473
Jorgensen, B. See Wild, C., 343 Kautsky, N., 400
Jorgensen, B.B. See Fossing, H., 316 See Chopin, T., 311
See Kuypers, M.M., 323 See Pihl, L., 403
See Schulz, H.N., 335 Kautsky, U. See Kautsky, N., 400
Jørgensen, B.B., See Boudreau, B.P., 126 Kawaguchi, K. See Takahashi, K., 136
Jorissen, F. See Barmawidjaja, D., 394 Kawahata, H. See Jickells, T.D., 465
Jorissen, F.J. See Pukaric, S.B.G.W., 403 Kay, M.C. See Lotze, H.K., 401
Juhl-Noodt, H., 82 Keatley, M.R., 465
Julian, P.R. See Madden, R.A., 325 Keddy, P.A., 400
Jumars, P.A., 89–138, 131 Keefer, D.K., 322
See Abello, H.U., 124 Keel, S. See Sayre, R., 335
See Karp-Boss, L., 35 Kelaher, B.P., 322
See Kringel, K., 131 Kelfkens, G., 465
See Mayer, L.M., 133 Kelleher, G., 400
See Penry, D.L., 134 Keller, A.A., 465
See Taylor, L.H., 137 Keller, B.D. See Dayton, P.K., 76
Jung N. See Bernstein, B.L., 75 See Dayton, P.K., 313
Junge, C.E., 34 Kelly, J.C. See Lie, U., 169
Kelly, P.J. See Codispoti, L.A., 311
Kemeny, J.G., 131
K Kemp, D.J., 465
Kemp, L. See Druehl, L.D., 78
Kaartvedt, S., 131 Kendall, M.A., 465
Kabat, P.W., 400 See Mieszkowska, N., 469
504
AUTHOR INDEX
Kendrick, G.A. See Goldberg, N.A., 462 Kirkman, H., 466

See Walker, D.I., 475 Kirkmann, H. See Gerard, V.A., 79
Kennedy, F. See Jaramillo, E., 321 Kirschtel, D. See Hillebrand, H., 34
Kennedy, H. See Hays, G.C., 130 Kirugara, D. See Wahl, M., 475
Kennedy, R.J., 400 Kita, J. See Kikkawa, T., 466
Kennelly, S.J., 465 Kitchen, J.C., 35
Kenner, M.C., 82 Kitching, R.L. See Boulter, S.L., 455
Kennish, M.J., 400 Kiyohara, M. See Gao, K., 461
Kenyon, R.A. See Vance, D.J., 475 Kjerfve, B.C.D.R. See Morris, J.T., 402
Keogh, J.A. See Brown, M.T., 75 Klaus, R. See Turner, J.R., 194
See Nyman, M.A., 84 Klein-MacPhee, G. See Keller, A.A., 465
Keogh, K.E. See Darling, J.D., 127 Kleppel, G.S., 466
Keough, M.J., 322, 466 Kleypas, J. See Hughes, T.P., 464
See Downes, B.J., 77 See Sarmiento, J.L., 473
Kerker, M., 35 Kleypas, J.A., 466
Kerr, J. See Vance, D., 475 Klingbeil, R. See Leet W.S., 82
Kerr, R.A., 322 Klinger, T., 322
Kerrigan, R.A. See Clarke, P.J., 457 See DeWreede, R.E., 77
Kerry, K.R. See Ross, G.J.B., 472 Klos, E. See Keller, A.A., 465
Kerville, S.P. See Campbell, S.J., 456 Klovaité, K. See Olenin, S., 402
Keser, M., 466 Klueter, A. See Wild, C., 343
Kesselmeier, J. See Wilhelm, C., 476 Klyashtorin, L. See Beamish, R.J., 454
Kester, D.R. 466 Knight, A.W. See Sitts, R.M., 136
Key, R. See Davidson, N.C., 396 Knight, T.M. See Sutherland, W.J., 338
Key, R.M. See McNeil, B.I., 469 Knottnerus, O.S., 400
See Orr, J.C., 470 See Lotze, H.K., 401
Khan, P.K., 192 Knowlton, N., 466
Khokiattiwong, S., 192 Knox, G.A., 322
Kibirige, I., 131 Knudsen, E. See Jonzén, N., 465
See Perissinotto, R., 134 Kobayashi, J.-I. See Yamamoto, M., 138
Kidwell, S. See Jackson, J.B.C., 399 Koch, E. See Menzel, A., 469
Kidwell, S.M. See Lotze, H.K., 401 Kocifaj, M., 35
Kiefer, D.A. See Stramski, D., 36 Koeve, W. See Antia, A.N., 454
Kienzle, M. See Beare, D.J., 454 Koh, B.S. See Guizien, K., 318
Kiflawi, M., 169 Kokhanovsky, A.A., 35
Kikkawa, T., 466 Komar, P.D., 35
Kim, J.Y. See Porteiro C., 471 See Baba, J., 33
Kim, K. See Harvell, C.D., 463 Kong, I. See Castilla, J.C., 310
Kim, K.-Y. See Suh, H.-L., 136 See Ortlieb, L., 330
Kim, S.L., 82, 131, 322 Konstant, W.R. See Brooks, T.M., 395
Kimmance, S. See Hiscock, K., 399 Köpcke, B., 131
Kimmerer, W.J., 131 Kopczak, C.D., 82
Kimura, R.S., 82 Korhs, D.G. See Zimmerman, R.C., 477
Kindscher, K. See Su, J.C., 171 Kornfield, I. See Cárdenas, L., 309
King, B.R., 466 See Poulin, E., 332
Kingsolver, J.G.,See Helmuth, B., 463 Kornicker, L.S. See Odum, H.T., 170
Kinlan, B.P., 82, Kornilovs, G. See Möllmann, C., 133
See Broitman B.R., 75 Korringa, P., 400
See Reed, D.C., 85 Kortum, G., 322
See Siegel, D.A., 336 Koshiishi, Y. See Hirota, Y., 130
Kinley, J. See Wahl, M., 475 Koslow, J.A., 466
Kinloch, J.E. See Friedel, M.H., 461 See Richardson, A.J., 471
Kinloch, M. See Ward, T.M., 476 See Thresher, R., 474
Kinzie, R.A. See Buddemeier, R.W., 191 Kosro, P.M. See Strub, P.T., 338
Kiørboe, T. See Visser, A.W., 137 Köster, F.W. See Möllmann, C., 133
Kirby, M.X. See Jackson, J.B.C., 399 Kotta, I., 131
See Lotze, H.K., 401 Kotta, J. See Kotta, I., 131
Kirby, R.R., 466 Kouassi, E., 131
Kirk, J.T.O., 35 Kouwenberg, J.H.M., 466
505
AUTHOR INDEX
Kouysoubas, D. See Karakassis, I., 169 See Castilla, J.C., 310

Kowalczyk, E.A. See Gordon, H.B., 462 See Vargas, C.A., 341
Kozyreff, G. See See van Dalfsen, J. A., 405 Lagos, S.L. See Dean, T.A., 77
Kraemer, G.P. See Chopin, T., 311 Lagos, V. See Collantes, G., 311
Krause-Jensen, D. See Boström, C., 395 Laidre, K.L., 82
Krautz, C. See Escribano, R., 314 LaJeunesse, T.C., 192
Krautz, M.C. See Hückstädt, L.A., 320 Lake, S., 132
Kreeger, D.A. See Kreeger, K.E., 131 Lakso, A.N. See Wolfe, D.W., 477
Kreeger, K.E., 131 Lambeck, K. See Church J.A., 457
Kremb, S. See Wild, C., 343 Lambshead, P.J.D, 169
Kremer, J.N. See Kopczak, C.D., 82 See Gage, J.,
See Zimmerman, R.C., 88 See Paterson, G.L.J., 170
Kremling, K. See Antia, A.N., 454 See Ugland, K. I., 172
Krenz, C. See Menge, B.A., 327 Lammers, M.O. See Benoit-Bird, K.J., 125
Kringel, K., 131 Lamont, P.A. See Levin, L.A., 324
Kroencke, I., 323 Lampitt, R.S. See Hirst, A.G., 464
Kruse, V.A. See Lesser, M.P., 467 Lancellotti, D. See Hinojosa, I., 320
Krylova, E. See Sellanes, J., 335 Lancellotti, D.A., 323
Kubilay, N. See Jickells, T.D., 465 See Stotz, W., 338
Kuffner, I.B., 466 See Thiel, M., 199–344
Kuhn, M. See Church J.A., 457 Landaeta, M.F., 323
Kuijpers, L. See de Jager, D., 458 See Castro, L.R., 311
Kullberg, J. See Parmesan, C., 470 Lande, R., 169
Kumar, H.D. See Häder, D.P., 462 Lane, C.D. See Druehl, L.D., 77
Kunz, T.J. See Hobday, A.J., 464 Lane, C.E., 82
See Poloczanska, E.S., 407–478, 466 Lanfranco, E. See Barbera, C., 394
Kunze, E., 131 See Boudouresque, C.F., 395
Kuosa, H. See Linden, E., 132 Langar, H. See Meinesz, A., 401
See Viherluoto, M., 137 Langdon, C., 466
Kura, Y. See Burke, L., 396 See Kleypas, J.A., 466
Kurashov, V. See Beamish, R.J., 454 Langdon, C.J. See Kreeger, K.E., 131
Kurihara, H., 466 Lange, C.B. See Escribano, R., 314
Kuss, J. See Antia, A.N., 454 See Fuenzalida, R., 316
Kuver, J. See Fossing, H., 316 See Jackson, J.B.C., 399
Kuypers, M.M., 323 See Milessi, A., 327
See Morales, C.E., 328
See Muñoz, P., 329
L Langenbuch, M. See Pörtner, H.O., 471
Langer, M. See Wahl, M., 475
Labanauskas, V. See Worm, B., 405 Langhanki, N. See Lotze, H.K., 401
Laborel, J. See Boudouresque, C.F., 395 Lankerani, A. See Hannah, L., 399
Labra, F.A. See Ojeda, F.P., 330 Lankford, T.E., 132
See Tognelli, M.F., 340 Laptikhovsky, V., 169
Lacis, A.A. See Mishchenko, M.I., 35 Lapyonok, T. See Dubovik, O., 34
Ladah, L.B., 82, 323 La Rocca, B. See Sfriso, A., 404
Ladds, P.W. See Adnyana, W., 453 Lardies, M.A., 323
Lafferty, K.D., 82 Largier, J. See Castilla, J.C., 310
See Airamè, S., 393 See Narváez, D.A., 329
See Behrens, M.D., 75 See Wieters, E.A., 343
See Roberts, C.M., 333 Largier, J.L., 323
See Ward, J.R., 476 See Botsford, L.W., 307
Laffoley, D. See Mieszkowska, N., 469 See Kaplan, D.M., 322
Laffoley, D.D., 400 See Shanks, A.L., 86
See Davidson, N.C., 396 See Wing, S.R., 88, 343
Lagardère. J.P., 131 Larichev, V.D. See Griffies, S.M., 462
Lagos, N. See Astorga, A., 306 Larkum, A.W.D., 466
See Correa, J.A., 312 See West, R.J., 476
Lagos, N.A., 323 LaRoche, J. See Boyd, P.W., 456
See Camus, P.A., 309 See Jickells, T.D., 465
506
AUTHOR INDEX
Larour, M. See Fabri, M.C., 167 Lefevre, N. See González, H.E., 317
Larson, R.J. See Stephens, J.S., Jr, 86 See Torres, R., 340
Larsson, C.S. See Lundälv, T., 401 Legakis, A. See Costello, M.J., 167
Larsson, U. See Hansson, S., 130 Legendre, L. See Goffart, A., 462
Lasenby, D. See Quirt, J., 134 Legendre, P., 169
Lasenby, D.C., 132 Leggett, M.F. See Johannsson, O.E., 130
Laslo, I. See Zhao, T.X.-P., 36 Leggett, W.C. See Choi, J.S., 126
Lassus, P. See Holmer, M., 399 See Frank, K.T., 129
Last, P.R. See Barrett, N.S., 397 Lehikoinen, A. See Jonzén, N., 465
Lastra, M. See Barbera, C., 394 Lehikoinen, E., 467
Latif, M. See Allan, R., 190 Lehodey, P., 467
Latimer, P., 35 Lehtiniemi, M. See Linden, E., 132
Laur, D.R. See Ebeling, A.W., 78 Leibold, M.A., 323
See Harris, L.G., 80 Leighton, D.L., 82
See Hempel, W.M., 80 Leising, A.W., 132
See Reed, D.C., 85 Leiva, F. See Bertrand, A., 307
Laval, B.E. See Hurd, C.L., 81 Leiva, G. See Narváez, D.A., 329
Lavery, P. See Lemmens, J., 323 See Poulin, E., 332
Lavik, G. See Kuypers, M.M., 323 Le Maho, Y. See Ferraroli, S., 460
Lavín, M.F. See Fiedler, P.C., 316 Lemaire, S. See Ene, A., 460
Lavoie, D.R., 82 Le Mao, P., 132
Law, C.S. See Boyd, P.W., 456 Lemmens, J., 323
Law, R. See Dieckmann, U., 128 Lenhian, H.S. See Jackson, J.B.C., 399
See Frid, C., 398 See Lotze, H.K., 401
See Leibold, M.A., 323 Lennon, J.J. See Thomas, C.D., 474
Lawrence, G.A. See Hurd, C.L., 81 Lenz, J. See Ene, A., 460
Lawrence, J.M., 82 See Herbst, L., 463
Lawrie, S. See Speirs, D., 136 León, R.I., 324
Lawrie, S.M., 132 Leonard, G.H., 82
See Thrush, S.F., 171 Le Pennect, M. See Moragat, D., 328
Lawson, G.L., 132 Lépez, I. See Acuña, E., 304
Lawton, J.H. See Jones, C.G., 322 Lépez, M.I. See Moreno, C.A., 328
Lea, D.W. See Hansen, J., 462 Leslie, H., 324, 400
See Swearer, S.E., 339 See Airamè, S., 393
See Zacherl, D.C., 344 See Menge, B.A., 327
Leaper, R. See Mieszkowska, N., 469 See Roberts, C.M., 333
See Simkanin, C., 473 Lesser, M.P., 467
Lear, D. See Mieszkowska, N., 469 See Shick, J.M., 473
Learmonth, J.A., 466 Lester, R.J.G. See Goggin, C.L., 462
See MacLeod, C.D., 468 Letelier, J., 324
See Robinson, R.A., 472 See Hormazábal, S., 320
Lebel, R.R. See Webster, P.J, 194 See Morales, C.E., 328
Leber, K.M., 323 Letelier, S., 324
Leblanc, K.W.G. See Syvitski, J.P.M., 36 Leterme, S.C. See Edwards, M.J., 459
Lechuga, A. See Hamm, L., 399 Leth, O., 324
Leconte, M. See David, V., 127 See Hormazábal, S., 320
Ledoyer, M., 132 Le Tissier, M.D.A. See Brown, B.E., 191
Lee, C.E., 323 See Scoffin, T.P., 193
Lee, H., II. See Behrenfeld, M., 455 Le Treut, H. See Bopp, L., 455
Lee, J.Y. See Yoon, H.S., 88 Levin, L. See Arntz, W.E., 306
Lee, K.K., 467 See Muñoz, P., 329
Lee, M., 400 See Wishner, K.F., 344
Lee, S. See Makris, N.C., 132 Levin, L.A., 169, 324
Lee Long, W.J. See Carruthers, T.J.B., 457 See DiBacco, C., 313
Leech, D.I. See Robinson, R.A., 472 See Gallardo, V.A., 316
Leet W.S., 82 See Gooday, A.J., 318
LeFaou, Y. See Glemaréc, M., 398 See Helly, J.J., 319
Le Fèvre, J., 323 See Tegner, M.J., 87
Lefevre, J.R. See Meinesz, A., 401 Levins, R., 169
507
AUTHOR INDEX
Levitus, S. See Da Silva, A.M., 312 Llanos, A., 324

Lewis, A. See Lehodey, P., 467 See Balbontín, F., 306
Lewis, C.V.W. See Miller, C.B., 133 See Castro, L.R., 311
Lewis, M. See Boss, E., 33 Llanos-Rivera, A., 324
Lewis, M.R. See Twardowski, M.S., 36 Lleellish, J., 324
Lewis, R., 82 Lleellish, M., 83
Lewis, R.J. See Reed, D.C., 85 Llewellyn, G. See Kaufman, L., 322
Li, N.K., 467 See Mumby, P.J., 469
Li, R., See Brawley, S.H., 456 Lluch-Cota, S.E. See Chávez, F.P., 311, 457
Lichtensztajn, Y.G. See Meltzoff, S.K., 327 Lo, K. See Hansen, J., 462
Liddicoat, M. See Boyd, P.W., 456 Lobban, C.S., 83
Lie, U., 169 See Barrales, H.L., 75
Lieberknecht, L.M. See Connor, D.W., 396 See Schmitz, K., 86
Light, K. See Graham, M.H., 79 Locarnini, R.A. See Conkright, M.E., 311
See Harrold, C., 80, 319 Lock, K., 132
Lillywhite, P. See MacNally, R., 169 Lockwood, D.R., 324
Limbaugh, C. See Feder, H.M., 78 Loeb, V.J., 324
Limburg, K. See Costanza, R., 457 Loeng, H. See Frid, C., 398
See Costanza, R., 396 Loew, E.R. See Losey, G.S., 467
Limpus, C.J., 467 Logan, B.W., 467
See Bowen, B.W., 455 Loh, W.K.W. See LaJeunesse, T.C., 192
See Loop, K.A., 467 Lohr, M. See Wilhelm, C., 476
See Marsh, H., 468 Lohrer, A. See Thrush, S.F., 474
Limtrakulvong, V. See Nielsen, T.G., 192 Lohrer, A.M., 467
Lin, B. See Hu, Y.-X., 34 Lohse, D. See Hannah, L., 399
Lin, H.-X. See van Dalfsen, J. A., 405 Loke Ming, C. See Hesp, P.A., 192
Lindberg, D.R., 83 Lombardo, M. See Trombini, C., 404
See Castilla, J.C., 310 Lomolino, M.V. See Brown, J.H., 308
Lindeboom, H.J., 401 Loneragan, N. See Abal, E.G., 453
Lindeman, K.C. See Mumby, P.J., 469 Loneragan, N.R., 467
Lindén, A. See Jonzén, N., 465 See Manson, F.J., 468
Linden, E., 132 See Vance, D.J., 475
Linderholm, H.W. 467 Long, D. See Roberts, J.M., 472
Lindley, J.A. See Beaugrand, G., 454 Long, S.P. See Ainsworth, E.A., 453
See Kirby, R.R., 466 Long, W.J.L. See Preen, A.R., 471
Lindsay, K. See Orr, J.C., 470 Longhurst, A.R., 132
Lindstrom, S.C. See Gabrielson, P.W., 79 Longstaff, B.J. See Carruthers, T.J.B., 457
Lindström M., 132 Lonhart, S.I. See Zacherl, D., 477
Ling, R. See Boyd, P.W., 456 Lonsdale, P.F. See Genin, A., 461
Ling, S. See Johnson, C., 465 Loop, K.A., 467
Linger, F.I. See Enbysk, B.J., 128 Lopatin, V.N. See Shepelevich, N.V., 36
Link, J.S., 132 López, C. See Stotz, W., 338
Linse, K. See Brandt, A., 126 López, C.A., 324
Lipp, E.K. See Harvell, C.D., 463 Lopez, M.D.G. See Huntley, M., 464
Lipschultz, F. See Codispoti, L.A., 311 Lopez Cazorla, A.C. See Sardiña, P., 135
Liqiang, H. See Rhoads, D.C., 471 López Gappa, J.J. See Orensanz, J.M., 134
Lisin, S. See Graham, M.H., 79 López-Urrutia, A. See Bonnet, D., 455
See Harrold, C., 80 Loreau, M. See Leibold, M.A., 323
See Harrold, C., 319 Loschnigg, J.P. See Webster, P.J, 194
Liss, P, See Raven, J., 471 Losey, G.S., 467
Liss, P.S. See Jickells, T.D., 465 Losos, E. See Wilcove, D.S., 405
Lissalde, J-P. See Cornet, M., 127 Lostaunau, N. See Codispoti, L.A., 311
Litchman, E., 467 Lothrop, S.K., 324
Little, C.M. See Donner, S.D., 459 Lotsberg, J.K. See Erga, S.R., 460
Little, M., 467 Lotze, H. See Schories, D., 403
Liu, P.C. See Lee, K.K., 467 Lotze, H.K., 401
Liu, S.L., Häder, D., 462 See Wahl, M., 475
Livingstone, D.R., 324 See Worm, B., 405, 477
Llagostera, A., 324 Lough, J. See Done, T.J., 459
508
AUTHOR INDEX
Lough, J.M., 192, 467 Macchiavello, J.E. See Bulboa, C.R., 308
See Hughes, T.P., 464 See Edding, M., 313
Loureau, M., 169 See Macaya, E.C., 83, 325
Loureiro, M. See Rodríguez-Graña, L., 333 MacDonald, B.A., 325
Love, M. See Stephens, J.S., Jr, 86 MacDonald, I. See Norkko, A., 470
Lovengreen, C. See Gómez, I., MacDonald, J.B. See Twardowski, M.S., 36
See Huovinen, P., 321 Mace, G.M., 169
Low, P.J. See Perry, A.L., 470 Machias, A. See Karakassis, I., 169
Lowe, J.A. See Gregory, J.M., 462 Maciolek, N.J. See Grassle, J.F., 168
Lowry, J.K. See Koslow, J.A., 466 Macke, A., 35
Lowry, L.F., 132 Mackenzie, C.L., Jr, 401
Lowry, R.R. See Kreeger, K.E., 131 Mackie, A.S.Y., 169
Lozan J.L., 401 Mackinson, S. See Okey, T.A., 470
Lubchenco, J., 169 Macko, S.A. See Mayer, L.M., 133
See Grantham, B.A., 318 MacLeod, C.D., 468
See Leslie, H., 324 See Learmonth, J.A., 466
See Menge, J., 327 See Robinson, R.A., 472
See Roberts, C.M., 333 MacNally, R., 169
Lucas, M.I. See Attwood, C.G., 74 Maconochie, J.R. See Friedel, M.H., 461
Lucic, D. See Bonnet, D., 455 Macpherson, E. See Martin, D., 401
Ludynia, K., 324 Macquart-Moulin, C., 132
Luikart, G. See Sih, A., 404 See Champalbert, G., 126
Lumb, C.M., 401 Madden, R.A., 325
Lumsden, L.F. See MacNally, R., 169 Madec, G. See Bopp, L., 455
Luna-Jorquera, G., 325 Madhaven, B.B., 192
See Ellenberg, U., 314 Madin, L.P. See Brown, H., 126
See Guerra, C., 318 Madronich, S., 325
See Ludynia, K., 324 Madsen, F.J., 325
See Marín, V.H, 326 Madsen, H. T. See van Dalfsen, J. A., 405
See Mattern, T., 326 Måge, F. See Menzel, A., 469
See Simeone, A., 337 Maggs, C.A. See Wilson, S., 405
See Thiel, M., 199–344 See Birkett, D.A., 395
See Weichler, T., 343 Magin, G. See Brooks, T.M., 395
Lundälv, T., 401 Magnusson, J.B. See Bokn, T.L., 395
Lundberg, B. See Boudouresque, C.F., 395 Magoulas, A. See Cárdenas, L., 309
Lunecke, K.M. See Moreno, C.A., 328 Magurran, A.E., 169
Lüning, K., 83, 325, 401 Mahan, L.C. See Dayton, P.K., 76
Lürling, M. See Schippers, P., 473 Mahmoudi, B. See Okey, T.A., 470
Luschi, P., 467 Mahnken, C. See Beamish, R.J., 454
Luther, H. See Kangas, P., 400 Mahowald, N. See Jickells, T.D., 465
Luther, M.E. See Potemra, J.T., 193 Mahyiddin, D. See Baird, A.H., 190
Luxoro, C. G. See Thiel, M., 199–344 Maidment, D. See Dunton, K., 313
Lyle, J.M., 468 Maier, I., 83
See Henry, G.W., 463 See Müller, D.G., 83
See Welsford, D.C., 476 See Westermeier, R., 343
Lynch, D.R. See Miller, C.B., 133 Maier-Reimer, E. See Orr, J.C., 470
Lyne, V. See Harris, G.P., 463 Majkowski, J., 468
Majluf, P., 325
Makino, H. See Yamamoto, M., 138
M Makris, N.C., 132
Maldonado, A., 325
Maasch, K.A. See Sandweiss, D.H., 334 Maldonado, M.T. See Boyd, P.W., 456
MacArthur, R.H., 169 Malinarich, A. See Sielfeld, W., 336
Macaya, E. See Hinojosa, I., 320 Manley, S.L. See Arnold, K.E., 74
See Thiel, M., 199–344 See North, W.J., 84
Macaya, E.C., 83, 325 Mann, G., 325
MacCallum, I., 35 Mann, K.H., 83
Macchiavello, J., 325 See Friesen, J.A., 129
See Edding, M., 313 See Gerard, V.A., 79
509
AUTHOR INDEX
Manning, M. See de Jager, D., 458 Marshall, N.J. See Losey, G.S., 467
Manriquez, P.H., See Thiel, M., 199–344 Marshall, P. See Hughes, T.P., 464
Manríquez, K. See Palma, A.T., 331 Martens, P. See Kelfkens, G., 465
Manríquez, P.H., 325 Martin, C. See Levin, L.A., 324
See Vargas, C.A., 341 Martin, D., 401
See Zacherl, D.C., 344 See Airoldi, L., 394
Manríquez, R. See Escribano, R., 314 Martin, J.H., 326
Manson, F.J., 468 Martin, M. See Anderson, B.S., 74
See Loneragan, N.R., 467 Martínez, D. See Stotz, W., 338
Mantua, N.J., 468 Martínez, E.A., 326
Manzanera, M. See van Dalfsen, J. A., 405 See Faugeron, S., 315
Mao, C. X. See Colwell, R.K., 167 Martínez, G., 326
Marbà, N., 401 Martinez, N.D. See Dunne, J.A., 128
See Diaz-Almela, E., 458 Martínez, P., 326
See Duarte, C.M., 459 See Poulin, E., 332
Marchand, M. See Nicholls, R.J., 402 See Marbà, N., 401
Marchant, M., 325 See Vargas, C.A., 341
See Coloma, C., 311 Martínez, V. See Ahumada, R., 305
See González, H.E., 317 Marty, G. See Delille, D., 313
See Hebbeln, D., 319 Marubini, F. See Langdon, C., 466
Marchesiello, P., 325 Mas, J. See Badalamenti, F., 394
Marchetti, R., 401 Mascia, M., 326
Marchinko, K.B., 468 Mason, C.F., 468
See Arsenault, D.J., 454 Masotti, I. See Rutllant, J.A., 334
Marchioretti, M. See Jaubert, J.M., 400 Massuti, C. See Delgado, O., 396
Marcogliese, D.J. See Jackson, C.J., 130 Massuti-Pascual, E. See Meinesz, A., 401
Marcos, C. See Salas, F., 171 Masuero, A. See Lagos, N., 323
Marcos, R. See Gutierrez, A., 80 Matano, R.P. See See Mesías, J.M., 327
See Gutierrez, A., 319 Matear, R. See Orr, J.C., 470
Marcos-Ramirez, R. See Aguilar-Rosas, L.E., 74 See Poloczanska, E.S., 407–478
Marcovaldi, M.A. See Godfrey, M.H., 462 See Sarmiento, J.L., 473
Marcus, N.H., 325 Matear, R.J. See McNeil, B.I., 469
Mardal, W. See Moller, A.P., 469 Matern, S.A. See Feyrer, F., 128
Margalef, R., 169 Matranga, V. See Bonaventura, R., 455
Marin, V. See Giraldo, A., 317 Matsuhiro, B. See Vásquez, J.A., 341, 342
See Moreno, C.A., 328 Matsumoto, G.I., 132
See Thiel, M., 199–344 Matsuoka, M. See Hanamura, Y., 130
Marín, V.H, 325, 326 Matsuura Y. See Porteiro C., 471
See Escribano, R., 314 Matsuzawa, Y. See Sato, K., 473
See González, A., 317 Mattern, T., 326
See Rojas, P.M., 333 See Ellenberg, U., 314
Marincovich, L., Jr, 326 Mattila, J., 132
Mark, F. See Brante, A., 308 Mattison, J.E., 83
Markkula, S.E., 468 Maucher, J.M. See Hutchins, D.A., 321
Marocco, R. See Bondesan, M., 395 Mauchline, J., 132
Marques, J.C. See Azeiteiro, U.M.M., 125 Maudire, G. See Fabri, M.C., 167
See Salas, F., 171 Maurer, D., 133Maurer, B.A., 169
Marquet, P. See Alvarado, J.L., 305 Mauri, M. See Simonini, R., 404
Marquet, P.A., 326 Maxwell, J.G.H., 468
See Fernández, M., 315 May, R.M., 169
See Santelices, B., 335 See Gage, J., 167
See Santoro, C.M., 335 Mayer, L.M., 133
See Tognelli, M.F., 340 Mayes, C. See Druehl, L.D., 78
See Valdovinos, C., 172, 340 See Lane, C.E., 82
Marra, P.P., 468 Mayhoub, H. See Boudouresque, C.F., 395
Marsac, F. See White, W.B., 476 Mayor, S. See Gutiérrez, D., 319
Marsaleix, P. See Guizien, K., 318 Mazza, J. See Brunet, M., 126
Marsh, H., 468 McAllan, I.A. See Beaumont, L.J., 454
Marsh, T.D. See Beck, M.W., 395 McAllister, D. See Burke, L., 396
510
AUTHOR INDEX
McArdle, D. See Pandolfi, J.M., 402 McPeak, R.H., 83

See Roberts, C.M., 333 See Aguilar-Rosas, L.E., 74
See Airamè, S., 393 See Barilotti, D.C., 75
McArthur, L.C., 468 McQuaid, C.D. See Attwood, C.G., 74
McCann, K., 133 See Kaehler, S., 82
See Rooney, N., 135 See Perissinotto, R., 84
McCann, K.S. See Vadeboncoeur, Y., 137 See Teske, P.R., 339
McClain, C.R. See Rex, M.A., 170 McQuoid, M., 469
McClanahan, T. See Defeo, O., 313 McRoy, C.P. See McConnaughey, T., 133
McClean, C.J. See Balmford, A., 307 McShane, P.E., 469
McCleery, R.H. See Votier, S.C., 475 McWilliams, J.C. See Marchesiello, P., 325
McClenachan, L. See Pandolfi, J.M., 402 See Shchepetkin, A.F., 336
McCleneghan, K., 83 Mecum, W. See Orsi, J.J., 134
McComb, A.J. See Walker, D.I., 475 Medina, M., 326
McConnaughey, T., 133 Medina, R. See Minello, T., 402
McConnico, L., 83 Medina-Elizade, M. See Hansen, J., 462
McCrary, M.D. See Dugan, J.E., 313 Medrano, S. See Carrasco, F.D., 309
McCulloch, A. See Shanks, A.L., 336 Meehl, G.A. See Cubasch, U., 458
McDarby, J. See See van Dalfsen, J. A., 405 Mees, J., 133
McDougall, K. See Mullins, H.T., 329 See Beyst, B., 125
McDougall, T.J. See Walsh, K.J.E., 476 See Dewicke, A., 128
McEvedy, C., 401 See Hostens, K., 130
McFarland, W.N. See Losey, G.S., 467 See Lock, K., 132
McFarlane, G.A. See Beamish, R.J., 454 Meesters, E.H. See Bak, R.P.M., 190
McGarvey, R. See Ward, T.M., 476 Meier, O.W. See Porter, J.W., 471
McGehee, D.E. See Greenlaw, C.F., 129 Meinesz, A., 401
McGill, H., 170 See Boudouresque, C.F., 395
McGowan, J. See Roemmich, D., 472 Meir, E., 327
McGowan, J.A., 468 Meland, K. See Brattegard, T., 126
See Haury, L.R., 319 Meltzoff, S.K., 327
See Venrick, E.L., 475 Mena, O. See Sánchez, M., 334
McGrath, D. See Simkanin, C., 473 Méndez, C.A., 327
McGregor, J.L. See Gordon, H.B., 462 Mendo, J. See Arntz, W.E., 306
McInnes, K. See Pittock, A.B., 470 See Pauly, D., 331
McInnes, K.L. See Walsh, K.J.E., 476 See Stotz, W., 338
McInnis, H. See Sandweiss, D.H., 334 See Wolff, M., 344
McKay, R.M. See Boyd, P.W., 456 Meneses, I., 327
McKee, D. See MacCallum, I., 35 See González, J., 318
McKenzie, E. See Beare, D.J., 454 See Santelices, B., 335
McKenzie, L.J. See Campbell, S.J., 456 See Vega, J.M.A., 342
See Carruthers, T.J.B., 457 Menge, B.A., 327
McKenzie, R.L., 468 See Connolly, S.R., 311
See Madronich, S., 325 See Grantham, B.A., 318
McKinnon, A.D. 468 See Leslie, H., 324
See Alongi, D.M., 453 See Russell, R., 171
McLachlan, A., 326 Menge, D.N.L. See Menge, B.A., 327
See Brown, A.C., 395 Menkhorst, P.W. See Ross, G.J.B., 472
See Defeo, O., 313 Menzel, A., 469
McLachlan, J. See Avila, M., 306 See Walther, G.R., 476
McLaren, I. See Escribano, R., 314 Menzies, R.J., 327
McLean, J.H., 83 Merino, A. See Collantes, G., 311
McLean, S.R. See Gaylord, B., 79 Merino, G.E. See von Brand, E., 343
McLeay, L. See Ward, T.M., 476 Mesías, J.M., 327
McLoughlin, K. See Caton, A.., 457 See Strub, P.T., 338
McMahon, C.R., 468 Messner, U., 401
McManus, G.B. See Peterson, W.T., 331 Mestre, A. See Menzel, A., 469
McManus, J.W., 192 Metcalfe, J.D. See Hays, G.C., 463
McMillan, C., 469 Metcalfe, N.B. See Both, C., 455
McNeil, B.I., 469 Metillo, E.B., 133
511
AUTHOR INDEX
Metz, J.A.J. See Dieckmann, U., 128 Moe, R.L., 83

Mews, M. See Orr, M., 84 Moenne, A. See Correa, J.A., 312
Meyer, C.A. See Okey, T.A., 470 Moffat, A.M., 133
Meyer, C.P. See Castilla, J.C., 310 Moffat, C. See Figueroa, D., 316
Meyer-Rochow, V.B. See Lindström M., 132 Moffitt, R.B. See Polovina, J.J., 471
Meyers, G. See Harris, G.P., 463 Mohammadian, M.A. See Johannsson, O.E., 130
Meza, J. See Simeone, A., 337 Mohan, B.K. See Madhaven, B.B., 192
Mianzan, H.W. See Schiariti, A., 135 Molinero, J.C. See Bonnet, D., 455
Michaelidis, B., 469 Molis, M. See Valdivia, N., 340
Michaels, M. See Simeone, A., 337 See Wahl, M., 475
Micheli, F., 327 Moller, A.P., 469
See Harborne, A.R., 463 Moller, I. See Wolters, M., 405
See Pandolfi, J.M., 470 Möller, P. See Westermeier, R., 88
Middelboe, A.L., 401 Möllmann, C., 133
Middelburg, J.J. See Dauwe, B., 127 Moloney, C.L., 327
See Duarte, C.M., 459 See Botsford, L.W., 307
Mie, G., 35 Monaghan, P. See Furness, R., 316
Mieszkowska, N., 469 Monfray, P. See Bopp, L., 455
See Simkanin, C., 473 See Orr, J.C., 470
Mikolajewicz, U. See Sarmiento, J.L., 473 See Sarmiento, J.L., 473
Milazzo, M., 402 Montanari, G. See Simonini, R., 404
Milessi, A., 327 Montecino, V., 327
Miller, C.B., 133
See Pizarro, G., 332
See Crain, J.A., 127
See Rutllant, J.A., 334
Miller, J. See Shanks, A.L., 336
See Strub, P.T., 338
Miller, J.D. See Loop, K.A., 467
Montecinos, A., 328
Miller, K. See Johnson, C., 465
Montenegro, C. See Quiroz, J.C., 332
See Roughgarden, J., 333
Montero, P. See Cuevas, L.A., 312
Miller, M.W. See Baums, I.B., 307
See Daneri, G., 312
Milliman, J.D., 469
See Troncoso, V.A., 340
Mills, E.L., 170
Montoya, J.M., 133
Milne, A. See Clarke, R.B., 166
Montu, M.A. See Gama, A.M. da S., 129
Milne, D.J. See Bowman, D.M.J.S., 456
Mooj, W.M. See Grimm, V., 129
Milton, D.A. See Poloczanska, E.S., 407–478
Moore, A.M. See Webster, P.J, 194
Mimura, N. See Burkett, V., 456
Minamikawa, S. See Sato, K., 473 Moore, C. See Latimer, P., 35
Mincks, S. See Smith, C.R., 337 Moore, D.R. See Thomas, L.P., 474
Minello, T., 402 Moore, F.R. See Marra, P.P., 468
Minello, T.J. See Beck, M.W., 394 Moore, J.C. See Rooney, N., 135
Minghelli-Roman, A. See Jaubert, J.M., 400 Moore, J.D. See Vilchis, L.I., 475
Miralto, A. See Ianora, A., 321 Moore, L.B., 83
Miranda, A. See Bonnet, D., 455 Moore, L.R., 469
Miranda, C., 327 Moore, M.V. See Smith, S.L., 136
Miranda-Esquivel, D.R. See Posadas, P., 170 Moore, P. See Mieszkowska, N., 469
Mishchenko, M.I., 35 Moore, P.G., 328
See Macke, A., 35 See Barbera, C., 394
See Dubovik, O., 34 See Donnan, D.W., 397
Mitchell, B.G., 469 See Hall-Spencer, J.M., 399
See Graham, M.H., 80 Moorthy, P., 469
Mitchell, C.E. See Harvell, C.D., 463 Mora, J. See Barbera, C., 394
Mitchell, R. See Davidson, N.C., 396 Moraga, J., 328
Mitchum, G.T. See Polovina, J.J., 471 See Acuña, E., 305
Mittermeier, C.G. See Brooks, T.M., 395 See Valle-Levinson, A., 341
Mittermeier, R.A. See Brooks, T.M., 395 See Weichler, T., 343
Mobley, C.D., 35 Moragat, D., 328
Mock, G., 402 Morales, C. See Bernal, P.A., 307
Modarressie, R., 469 See Grünewald A., 318
512
AUTHOR INDEX
Morales, C.E., 328 Moy, C.M., 329

See Böttjer, D., 307 Moy, F.E. See Bokn, T.L., 395
See Escribano, R., 314 Moyano, H., 329
See Hidalgo, P., 320 See Castilla, J.C., 310
Morales, E.G., 328 See Desqueyroux, R., 313
Morales, V. See Gómez, I., 317 Moyle, P.B. See Feyrer, F., 128
See Huovinen, P., 321 MPA Global, 402
Moran, S., 133 Mrosovsky, N., 469
Moreira, M.H. See Cunha, M.R., 127 See Glen, F., 461
Morel, A., 35 See Godfrey, M.H., 461 462
See Bricaud, A., 33 See Yntema, C.L., 477
See Stramski, D., 36 Muck, P. See Pauly, D., 331
Morel, F.M.M. See Palenik, B., 470 Mueller, E. See Downs, C.A., 191
See Rich, H.W., 471 Mueller, H. See Furukawa, K., 461
See Riebesell, U., 472 Muinonen, K. See Macke, A., 35
See Tortell, P.D., 475 Muir, P.R. See Wallace, C.C., 194
Moreno, C. See Buschmann, A.H., 308 Mujica, A. See Acuña, E., 304
Moreno, C.A., 83, 328 Mukminin, A. See Baird, A.H., 190
See Buschmann, A.H., 75 Mulkins, L.M., 133
See Castilla, J.C., 76 Müller, A.M. See Wilhelm, C., 476
See Duarte, W.E., 313 Müller, D.G., 83
See Fernández, M., 315 See Maier, I., 83
See Jara, F., 321 See Westermeier, R., 343
See Jara, H.F., 321 Müller, H. See Müller, D.G., 83
See Marín, V.H., 326 Mullineaux, L. See Wishner, K.F., 344
See Zamorano, J.H., 344 Mullins, H.T., 329
Moreno, R.A., 328, 329 Mulvihill, R.S. See Marra, P.P., 468
See Hernández, C.E., 319 Mumby, P.J., 469
See Sepúlveda, R., 336 See Harborne, A.R., 463
Moretti, R. See Ene, A., 460 Munda, I.M., 402
Morgan, L. See Guinotte, J.M., 462 Munday, B.W. See Ball, B.J., 394
Morgan, L.E. See Wing, S.R., 343 Munilla, T. See San Vicente, C., 135
Morgan, S.G. See Swearer, S.E., 339 Muñiz, L. See Ramorino, L., 333
Morgigni, M. See Sandulli, R., 403 Munk, P. See Nielsen, T.G., 192
Morri, C. See Sandulli, R., 403 Muñoz, A.A. See Ojeda, F.P., 330
Morris, D.W., 133 Muñoz, C. See Palma, A.T., 331
Morris, J.T., 402 Muñoz, G. See Hernández, C.E., 319
Morris, P.M. See Stephens, J.S., Jr, 86 Muñoz, I., 329
Morrison, W. See Thessen, A.E., 474 Muñoz, J. See Simeone, A., 337
Morrow, J.H. See Jaubert, J.M., 400 Muñoz, M., 329
Mortensen, P.B. See Roberts, J.M., 472 Muñoz, O. See Hoovenier, J.W., 34
Moschella, P.S., 402 Muñoz, P., 329
See Airoldi, L., 394 See Avaria S., 306
See Benedetti-Cecchi, L., 395 See Thiel, M., 199–344
See Jonsson, P.R., 465 Munoz, V., 83, 329
See Martin, D., 401 See Gutierrez, A., 80, 319
See Mieszkowska, N., 469 Munoz-Schick, M. See Castro, S.A., 311
Moseley, M.E. See Keefer, D.K., 322 Murano, M. See Hanamura, Y., 130
Moss, D. See Davies, C.E., 396 Murawski, S.A. See Fogarty, M.J., 129
Mouchet, A. See Orr, J.C., 470 Murdoch, R. See Boyd, P.W., 456
Mouchot, M.C. See Belsher, T., 75 Murphy, G.I. See Majkowski, J., 468
Mouny, P., 133 Murphy, J.J. See Lyle, J.M., 468
See Dauvin, J.-C., 127 Murray, J., 170, 329
See Zouhiri, S., 138 Murray, S.N. See Bokn, T.L., 395
Mouquet, N. See Leibold, M.A., 323 See Fain, S.R., 78
Mouritsen, K.N., 469 Murty, V.S.N. See Varkey, M.J., 194
Moverley, J. See Edgar, G.J., 459 Mustapha, K.B. See Meinesz, A., 401
513
AUTHOR INDEX
Myers, A. See Simkanin, C., 473 Neill, P.E., 329

Myers, R.A. See Worm, B., 477 See Castilla, J.C., 310
Myers, R.F., 192 Neira, C., 329
Myerscough, P.J. See Clarke, P.J., 457 See Arntz, W.E., 306
See Gutiérrez, D., 319
See Muñoz, P., 329
N See Sellanes, J., 335, 336
Neira, S., 330
Naeem, S. See Costanza, R., 396, 457 See Arancibia, H., 305
Naganuma, N. See Hirota, Y., 130 See Moloney, C.L., 327
Nagelkerken, I. See Dorenbosch, M., 459 Nejstgaard, J. See Engel, A., 460
Nagy, K.L. See Bickmore, B.R., 33 Nelson, C.S. See Bakun, A., 306
Naito, Y. See Sato, K., 473 Nelson, D.J. See Friedel, M.H., 461
Najjar, R.G. See Orr, J.C., 470 Nelson, P.A. See Losey, G.S., 467
Nakanishi, A. See Arana, P., 305
Nemerson, D.M., 133
Narayanaswamy, B.E., 170
Neori, A. See Chopin, T., 311
Narváez, D. See Poulin, E., 332
Nero, R.W. See Makris, N.C., 132
Narváez, D.A., 329
Nerzic, R. See Hyder, P., 192
See Piñones, A., 332
Neshyba, S. See Silva, N., 337
See Vargas, C.A., 341
Neuer, S. See Antia, A.N., 454
Narvarte, M.E., 329
Neushul, M., 83
Nash, R.D.M. See Oh, C.W., 133
See Amsler, C.D., 74
Nash, S.B. See Duke, N.C., 459
See Dawson, E.Y., 76
Nast, J. See Greve, W., 462
See Lewis, R., 82
National Research Council, 170
See Lüning, K., 83
Navanete, S.S. See Valdovinos, C., 172
See Nyman, M.A., 84
Navarrete, S. See Castilla, J.C., 310
See Reed, D.C., 85
See Kaplan, D.M., 322
Neville, C.M. See Beamish, R.J., 454
Newberger, P.A. See Enfield, D.B., 314
Navarrete, S.A., 329
See Aiken, C., 305 Newell, R.C., 402
See Broitman, B.R., 308 Newland, C. See Haury, L., 130
See Fernández, M., 315 Newman, M.J.H. See Pandolfi, J.M., 402
See Lagos, N.A., 323 Newman, W.A. See Hinojosa, I., 320
See Manríquez, P.H., 325 Nhuan, M.T. See Church J.A., 457
See Martínez, P., 326 Ni, J.F. See Guzman-Speziale, M., 191
See Menge, B.A., 327 Nicholls, N. See Allan, R., 190
See Narváez, D.A., 329 See Limpus, C.J., 467
See Neill, P.E., 329 Nicholls, P.E. See Thrush, S.F., 475
See Nielsen, K.J., 330 Nicholls, R.J., 402
See Piñones, A., 332 Nichols, P.D. See Thresher, R.E.,
See Poulin, E., 332 Nicholson, N.L., 84
See Thiel, M., 199–344 Nickell, T.D. See Narayanaswamy, B.E., 170
See Valdovinos, C., 340 Nielsen, J. See Jernakoff, P., 465
See Vargas, C.A., 341 Nielsen, K.J., 330
See Velázquez, I., 342 See Grantham, B.A., 318
See Wieters, E.A., 343 See Wieters, E.A., 343
Navarro, J. See Jaramillo, E., 322 Nielsen, L.P. See Fossing, H., 316
Navarro, J.C., 133 Nielsen, T.G., 192
Navarro, J.M. See Gajardo, G., 316 Niemi, Å. See Kangas, P., 400
Navea, M. See Alarcón, E., 305 Nietch, C.T. See Morris, J.T., 402
Neale, P.J. See Litchman, E., 467 Nieto, K. See Silva, C., 337
Neckles, H.A. See Short, F.T., 473 See Yáñez, E., 344
Neefus, C. See Chopin, T., 311 Nilsen, K.J. See Rhoads, D.C., 471
Neff, J.M., 133 Nilsson, C. See Harris, G., 463
Negri, A., 469 Nilsson, H.L. See Attramadai, Y.G., 125
Nehring, D., 402 See Lindström M., 132
Neil, D.M., 133 Nilsson, J., 402
Neill, P.A. See Moreno, R.A., 329 See Choo, K.S., 457
514
AUTHOR INDEX
Niquen, M. See Alheit, J., 305 O’Farrell, S.P. See Gordon, H.B., 462
See Chavez, F.P., 311, 457 See Gregory, J.M., 462
See Porteiro C., 471 Ogden, J. See Boersma, D., 307
Nisbet, R.M., 84 Ogden, J.C. See Pandolfi, J.M., 470
Nivet, C., 402 Oh, C.W., 133
Noakes, D.J. See Beamish, R.J., 454 See Hong, S.Y., 130
Noda, A. See Cubasch, U., 458 O’Hara, T. See Koslow, J.A., 466
Nodder, S. See Boyd, P.W., 456 O’Hara, T.D., 470
Noguchi, G.E. See Evans, M.S., 128 Ohme, U. See Santelices, B., 335
Norall, T. See Grove, R.S., 462 Ohtsuka, S., 134
Norambuena, R. See Santelices, B., 335 Oikari, A.O.J. See Huovinen, P.J., 81
Nordli, Ø. See Menzel, A., 469 Ojeda, B. See Sandweiss, D.H., 334
Norici, A. See Giordano, M., 461 Ojeda, F.P., 84, 330
Norkko, A., 470 See Angel, A., 305
See Thrush, S.F., 474 475 See Cárdenas, L., 309
Norkko, J. See Norkko, A., 470 See Fariña, J.M., 315
Norman, F.I. See Bunce, A., 456 See Fernández, M., 315
See Ross, G.J.B., 472 See Hernández-Miranda, E., 319
Norse, E.A., 170 See Palma, A.T., 330, 331
See Watling, L., 405 See Poulin, E., 332
North, W.J., 84 See Pulgar, J.M., 332
See Anderson, E.K., 74 See Santelices, B., 85, 335
See Gonzalez-Fragoso, J., 79 See Vásquez, J.A., 87, 341
See Wheeler, P.A., 88 Okaji, K. See Brodie, J., 456
Northen, K.O. See Connor, D.W., 396 Okey, T.A. See Hobday, A.J., 464
Norton, T.A., 84 See Poloczanska, E.S., 407–478, 470
Nothig, E.M. See González, H.E., 318 Oldfield, S. See Brooks, T.M., 395
Nouvel, H. See Lagardère, J-P., 131 Oldham, R.D., 192
Novitsky, J.A. See Friesen, J.A., 129 Olenin, S., 402
Nowell, A.R.M. See Jumars, P.A., 131 See Reise, K., 403
Nozais, C. See Kibirige, I., 131 Olesen, B. See Marbà, N., 401
See Perissinotto, R., 134 Oliger, P., 330
NRC, 330 Oliva, D. See Bernal, P.A., 307
Nultsch, W. See Hanelt, D., 462 Olivares, C. See Radashevsky, V.I., 332
Nuñez, L., 84, 330 Olivares, G., 330
See Briones, L., 308 See Escribano, R., 314
Nunez, M. See Harris, G.P., 463 See Marín, V.H., 326
Núñez, S. See Letelier, J., 324 Olivares, G.R. See Marín, V.H., 326
Núñez, S.P. See Arcos., 305 Olivares, J. See Valle-Levinson, A., 341
See Vargas, C.A., 341 Oliver, J., 470
Nusetti, O.A. See Brokordt, K.B., 308 Oliver, J.K. See Babcock, R.C., 454
Nyholm, N.E.I. See Both, C., 455 Oliver, J.S. See Kim, S.L., 131
Nyman, M.A., 84 Oliver, P.G. See Mackie, A.S.Y., 169
See Brown, M.T., 75 Olivieri, R.A., 330
Nystrom, M. See Hughes, T.P., 464 Olkin, I. See Hedges, L.V., 168
Olsgard, F., 170
See Somerfield, P.J., 171
O Olyarnik, S.V. See Byrnes, J., 75
Omori, M., 134
Oakley, J. See Hiscock, K., 399 Onbe, T. See Ohtsuka, S., 134
Obenat, S. See Orensanz, J.M., 134 O’Neal, J.P. See Mattila, J., 132
O’Brien, C.M., 470 O’Neill, R.V. See Costanza, R., 396, 457
O’Brien, J.J. See Potemra, J.T., 193 Ong, J.E. See Ewel, K.C., 460
O’Brien, T.D. See Conkright, M.E., 311 Opdyke, B.N. See Kleypas, J.A., 466
Occhipinti-Ambrogi, A., 402 Oppenheimer, M. See Donner, S.D., 459
Ocean Studies Board, 402 Ord, D.C. See Hempel, W.M., 80
Ochieng, C.A., 470 Orel, G. See Caressa, S., 396
Odaya, M. See Beck, M.W., 307, 395 Orensanz, J.M., 134
Odum, H.T., 170 See González, J., 318
515
AUTHOR INDEX
Orestano, C. See Meinesz, A., 401 See Castilla, J.C., 310

Orfila, A. See Marbà, N., 401 See Sebens, K.P., 335
Orhon, D. See Jeftic, L., 400 Painting, S.J. See Verheye, H.M., 342
O’Riordan, R. See Simkanin, C., 473 Pak, H. See Carder, K.L., 33
Orostegui, M. See Gómez, I., 317 See Kitchen, J.C., 35
Orr, J. See Guinotte, J.M., 462 See Zaneveld, J.R.V., 36
Orr, J.C., 470 Pakhomov, E.A. See Kaehler, S., 82
See Bopp, L., 455 Pal, T., 192
Orr, M., 84 Paladino, F.V. See Binckley, C.A., 455
Orrego, P. See Edding, M., 313, 314 Palenik, B., 470
Orsi, J.J., 134 Paleras, A. See Michaelidis, B., 469
See Kimmerer, W.J., 131 Palma, A.T., 330, 331
Orth, R.J. See Beck, M.W., 394 See Fariña, J.M., 315
Ortiz, V. See Clarke, M., 311 See Hernández-Miranda, E., 319
Ortíz, E. See Takesue, R.K., 339 See Poulin, E., 332
Ortíz, M., 330 Palma, F. See Sielfeld, W., 336
Ortíz, V. See Castilla, J.C., 310 Palma, M., 331
Ortíz, V.C. See González, H.E., 318 See Gallardo, V.A., 316
Ortlieb, L., 330 See Muñoz, P., 329
See Takesue, R.K., 339 Palma, S., 331
See Valdés, J., 340 Palma, W., 331
Osborn, J.E. See Foster, E.G., 129 See Pizarro, J., 332
Osborne, A.R., 192 Palmer, A.R., 331
Oshel, P.E. See Janssen, J., 130 See Arsenault, D.J., 454
Osorio, P. See Buschmann, A.H., 75, 308 See Marchinko, K.B., 468
OSPAR Commission, 402 Palomera, I. See Tudela, S., 137
Osses, J. See Thomas, A.C., 339 Palumbi, S.R., 331
Ostergaard, J.B. See Nielsen, T.G., 192 See Hughes, T.P., 464
Osterhaus, A.D.M.E. See Harvell, C.D., 463 See Lubchenco J., 169
Ostfeld, R.S. See Harvell, C.D., 463 See Sotka, E.E., 337
Otaiza, R. See Santelices, B., 85 Paluszkiewicz, T. See Huyer, A., 321
Otsuki, Y. See Wolfe, D.W., 477 Panayotidis, O. See Boudouresque, C.F., 395
Ouzounis, C. See Michaelidis, B., 469 Pandolfi, J.M., 402, 470
Overstreet, R.M. See Harvell, C.D., 463 See Hughes, T.P., 464
Oviatt, C. See Keller, A.A., 465 See Jackson, J.B.C., 399
Oxnevad, S. See Berge, J.A., 455 Pankajakshan, T. See Desai, B.N., 191
Oyarce, E. See Marín, V.H., 326 Pannacciulli, F. See Benedetti-Cecchi, L., 395
Oyarzún, C., 330 Pantoja, S., 331
See Chong, J., 311 See Bernal, P.A., 307
See Galleguillos, R., 316 See González, H.E., 318
See Muñoz, P., 329
Panutrakul, S., 192
P Paolini, P. See Lagos, N., 323
Papaconstantinou, C., 134
Pacanowski, R.C. See Griffies, S.M., 462 Papadopoulou, K.N. See Karakassis, I., 169
Pacheco, A., 330 Papenfuss, G.F., 84
Pacheco, C. See Alvarado, J.L., 305 Papi, F. See Luschi, P., 467
See Castilla, J.C., 310 Papi, I. See Piazzi, L., 403
Pacheco, R. See Poulin, E., 332 Pappalardo, P. See Fernández, M., 315
Packard, T.T. See Codispoti, L.A., 311 Paradis, G. See Zacherl, D.C., 344
Paddack, M.J., 84 Paramonov, L.E., 35
Padilla, D.K. See Klinger, T., 322 Paramor, O.A.L. See Hughes, R.G., 399
Pagano, M. See Kouassi, E., 131 Paranjape, M.A. See Conover, R.J., 127
Page, G. See Galbraith, H., 461 Pardal, M.A. See Salas, F., 171
Page, H.M. See Bomkamp, R.E., 307 Pardede, S.T. See Baird, A.H., 190
Pagès, F., 330 Pardi, G. See Piazzi, L., 403
See González, H.E., 317, 318 Pardo, L.M. See Fernández, M., 316
Paine, J. See Green, M.J.B., 398 See Palma, A.T., 331
Paine, R.T., 330 See Vásquez, J.A., 87, 341, 342
516
AUTHOR INDEX
Paredes, C. See Tarazona, J., 339 Pedersen, T.F. See Cowie, G.L., 312
See Pandolfi, J.M., 402 Pederson, D.K. See Duke, N.C., 459
See Sala, E., 334 Pedraza, M. See Landaeta, M.F., 323
Paredes, R. See Sandweiss, D.H., 334 Peelaers, D. See Remerie, T., 134
Paris, C.B. See Cowen, R.K., 457 Peet, R.K., 170
Park, G.S. See Takeuchi, T., 136 Pegau, W.S. See Boss, E., 33
Park, R. See Galbraith, H., 461 Pegau, W.S. See Twardowski, M.S., 36
Parker, D.M. See Polovina, J.J., 471 Peinert, R. See Antia, A.N., 454
Parker, D.O. See Davis, G., 458 Peirano, A. See Bianchi, C.N., 395
Parkinson, R.W., 470 See Meinesz, A., 401
Parks, J.E. See Pomeroy, R.S., 332 See Sandulli, R., 403
Parma, A.M. See González, J., 318 Peletier, M.A. See See van Dalfsen, J. A., 405
Parmenter, C.J. See Hewavisenthi, S., 463 Pellatt, J. See Votier, S.C., 475
Parmesan, C., 470 Peloquin, J. See Cordi, B., 457
See Walther, G.R., 476 Pemberton, D., 470
Parnell, P.E. See Dayton, P.K., 77, 458 See Ross, G.J.B., 472
See Seymour, R.J., 86 Peña, J. See Moragat, D., 328
Parra, I. See Sala, E., 334 Peña, T.S., 331
Parrish, C.C. See Richoux, N.B., 135 Peñaloza, M. See Galleguillos, R., 316
Parrish, F.A. See Polovina, J.J., 471 Peñalver, E., 331
Parsons, M.L. See Thessen, A.E., 474 Penchaszadeh, P. See Orensanz, J.M., 134
Paruelo, J. See Costanza, R., 396, 457 Pennington, J.T. See Roughgarden, J., 333
Pascual, M. See Orensanz, J.M., 134 Penry, D.L., 134
Pastorino, G. See Orensanz, J.M., 134 Peñuelas, J. See Menzel, A., 469
Pasqualini V., 402 Penven, P., 331
Paterson, D.M. See Lawrie, S.M., 132 Peredo, R. See Sielfeld, W., 336
Paterson, G.L.J., 170 Pereira, P. See Little, M., 467
Patino, D. See Westermeier, R., 343 Pere-Neto P.R. See Legendre, P., 169
Patrico, J. See Salas, F., 171 Perez, E. See Thiel, M., 199–344
Patriti, G. See Macquart-Moulin, C., 132 Pérez, E.P., 331
Patterson, H.M., 134 See Stotz, W., 338
Pauchard, A., 331 Pérez, M. See Invers, O., 464
Paulmier, A. See Cornejo, M., 312 Perez, T.,
Pauly, D., 331 See Garrabou, J., 461
Pávez, A. See González, H.E., 317 Perez-Llorens, J.L.See Bischof, K., 307
Pavéz, M. See Vargas, C.A., 341 Perez-Ruzafa, A. See Salas, F., 171
Pavez, M.A., 331 Pergent G. See Pasqualini V., 402
Pawlak, J.F. See Frid, C., 398 Pergent-Martini C. See Pasqualini V., 402
Pawson, D. See Levin, L.A., 169, 324 Perissinotto, R., 84, 134
Payá, I. See Santelices, B., 335 See Kibirige, I., 131
Peach, K. See Beare, D.J., 454 See Webb, P., 137
Pearce, A. See Caputi, N., 456 Perry, A.L., 470
Pearce, A.F. See Griffin, D.A., 462 Perry, C.J. See Abal, E.G., 453
Peard, K.R. See Weeks, S.J., 343 Perry, C.T. See Harborne, A.R., 463
Pearre, S., Jr, 134 Persson, G. See Rosenberg, R., 403
Pearse, J.S., 84 Persson, L.E. See Nilsson, J., 402
See Harrold, C., 80, 319 Pescod, C.L. See Mumby, P.J., 469
See Mattison, J.E., 83 Peters, A.F., 331
See Towle, D.W., 87 Peters, D. See Edgar, G.J., 459
Pearson, D.L., 170 Peters, R.H., 470
Pearson, G. See Serrão, E.A., 473 Petersen, C.G.J., 170
Pearson, G.A. See Brawley, S.H., 456 Petersen, K.S., 403
Pearson, T.H. See Tyson, R.V., 340 Peterson, C. See Kaufman L., 322
Pechenik, J.A., 331 Peterson, C.H. See Jackson, J.B.C., 399
Pedersén, M., 402 See Lotze, H.K., 401
See Choo, K.S., 457 Peterson, W.T., 331
See Johansson, G., 400 See Halpin, P.M., 319
Pedersen, M.F., 470 Petrie, B. See Frank, K.T., 129
Pedersen, S.E. See Rask, N., 403 Petrillo, M. See Thurston, M., 340
517
AUTHOR INDEX
Petryashev, V.V., 134 See Walsh, K., 475

Pettersen, O. See Berge, J.A., 455 See Walsh, K.J.E., 476
Petty, R.L. See Reed, D.C., 85 Pizarro, A., 332
Pezzoli, G. See Colombini, I., 311 Pizarro, G. See González, H.E., 317
Pfister, C.A., 84 See Iriarte, J.L., 321
Philippart, C.J.M., 470 See Montecino, V., 327
Phillips, A. See Wilcove, D.S., 405 Pizarro, J., 332
Phillips, J.A., 470 Pizarro, O. See Chaigneau, A., 311
Phillips, N.E., 332 See Escribano, R., 314
Phillips, R.A., 332 See Letelier, J., 324
Phillips, S. See Downs, C.A., 191 See Montecinos, A., 328
Phongsuwan, N., 193 See Ramos, M., 333
See Brown, B.E., 191 See Shaffer, G., 336
See Chansang, H., 191 Pizay, M.D. See Engel, A., 460
See Satapoomin, U., 193 Planque, B. See O’Brien, C.M., 470
Piacentini, D. See Jonzén, N., 465 Planqué, R. See See van Dalfsen, J. A., 405
Piazzi, L., 403 Plante, C.J. See Mayer, L.M., 133
Picaut, J. See Lehodey, P., 467 Plattner, G.K. See Orr, J.C., 470
Pickmere, S. See Boyd, P.W., 456 Pleijel, F. See Bertrand, Y., 166
Piel, M.I. See Westermeier, R., 343 Plenge, M. See Duffy, D., 313
Pielou, E.C., 170 Plummer, K.M., 134
Pienkowski, M.W. See Davidson, N.C., 396 Pohnert, G. See Ianora, A., 321
Pieper, R.E. See Smith, S.L., 136 Poiner, I. See Rothlisberg, P., 472
Pierce, G.J. See Learmonth, J.A., 466 Polacheck, T., 471
See MacLeod, C.D., 468 Polis, G.A., 332
See Robinson, R.A., 472 See Sanchez-Pinero, F., 334
Piersma, T. See Both, C., 455 Pollingher, U. See Hillebrand, H., 34
Pierson, M.O. See Dugan, J.E., 313 Polo, L. See Rodríguez-Prieto, C., 403
Pietras, C. See Zhao, T.X.-P., 36 Poloczanska, E.S., 407–478
Pihl, L., 134, 403 See Hobday, A.J., 464
See Baden, S., 394 See Mieszkowska, N., 469
Pikitch, E.K. See Boersma, D., 307 Polovina, J.J., 471
Pile, A.J. See Prowse, T.A.A., 471 Poma, V. See Bonaventura, R., 455
Pilgrim, J. See Brooks, T.M., 395 Pomeroy, R.S., 332
Pillai, C.S.G., 193 Ponce, F. See Morales, C.E., 328
Pillai, M.C. See Garman, G.D., 79 Ponce, T. See Simeone, A., 337
Piller, W.E. See Harrold, C., 319 Pondella, D.J., Jr. See Stephens, J.S., Jr, 86
Pilon, V. See Boulding, E.G., 455 Pool, R.M. See Wolfe, D.W., 477
Pimm, S.L. See Montoya, J.M., 133 Poore, G.C.B., 170, 471
Pineda, J., 170, 332 See Koslow, J.A., 466
See Levin, L.A., 169, 324 See O’Hara, T.D., 470
Pinedo, S. See Thibaut, T., 404 See Richardson, A.J., 471
Pinero, F.S. See Polis, G.A., 332 Poovachiranon, S. See Chansang, H., 191
Pinnick, R.G. See Ch∆lek, P., 34 Pope, N. See Roast, S.D., 135
Pino, M. See Jaramillo, E., 322 Popper, N. See Stambler, N., 194
Piñones, A., 332 Porflitt, G. See Guerra, C., 318
See Vargas, C.A., 341 Porteiro C., 471
Pinto, L. See Ahumada, R., 305 Porter, J.W., 471
Pinto, R. See Alvarado, J.L., 305 See Harvell, C.D., 463
See Martínez, E.A., 326 Pörtner, H.O., 471
Pipitone, C. See Badalamenti, F., 394 See Brante, A., 308
Piquet, A. See Kelfkens, G., 465 See Fernández, M., 316
Piraino, S. See Fanelli, G., 398 See Frederich, M., 316
Pirazzoli, P.A. See Bondesan, M., 395 See Michaelidis, B., 469
Pirinen, P. See Menzel, A., 469 See Ruiz-Tagle, N., 334
Piriz M.L. See Orensanz, J.M., 134 Posadas, P., 170
Pita, M.E. See Barbera, C., 394 Possingham, H., 403
Pitta, P. See Karakassis, I., 169 See Leslie, H., 400
Pittock, A.B., 470 See Roughgarden, J., 85,
518
AUTHOR INDEX
See Leslie, H., 324 Pye, K. See Allen, J.R.L., 394

See Meir, E., 327 Pyle, B.E. See Latimer, P., 35
See Pandolfi, J.M., 470
Post, E. See Walther, G.R., 476
Potemra, J.T., 193 Q
Potter, D.S. See Spencer, C.N., 136 Qin, D. See Church J.A., 457
Potti, J. See Both, C., 455 Quéguiner, B. See Sedwick, P.N., 473
Potts, D.C., 193 Quijón, P., 332
Pou, S. See Delgado, O., 396 See Contreras, H., 312
Poulet, S.A. See Ianora, A., 321 See Jaramillo, E., 322
See Vargas, C.A., 341 Quijón, P.A., 170
Poulin, E., 332 Quinby-Hunt, M.S., 36
See Cárdenas, L., 309 Quinn, J.F. See Botsford, L.W., 307
See Narváez, D.A., 329 See Wing, S.R., 343
See Thiel, M., 199–344 Quiñones, R. See Daneri, G., 312
Poulin, R. See Mouritsen, K.N., 469 See González, H.E., 317
Pounds, J.A. See Root, T.L., 472 Quiñones, R.A. See Cury, P., 312
Powell, H.T., 84 See Eissler, Y., 314
Powell, T.M. See Botsford, L.W., 307 See González, R.R., 318
Power, A. See Simkanin, C., 473 See Ulloa, O., 340
Pozdnyakov, D., 35 See Yaikin, J., 344
Pradel, R. See Rivalan, P., 472 Quirantes, A., 36
Prado, L., 332 Quiroga, E., 332
Prakash, A. See Sheldon, R.W., 36 See Palma, M., 331
Preen, A.R., 471 See Sellanes, J., 336
Preston, F.W., 170 Quiroz, D. See Báez, P., 306
Preti, M. See Simonini, R., 404 See Montecino, V., 327
Prevedelli, D. See Simonini, R., 404 Quiroz, J.C., 332
Prevost, C. See Delille, B., 77 Quirt, J., 134
Prévot-Julliard, A.C. See Rivalan, P., 472
Price, D.N. See Cordi, B., 457
Price, J.T. See Root, T.L., 472 R
Price, K.S. Jr, 134 Rabinowitz, C. See Baruch, R., 454
Price, N.M. See Palenik, B., 470 Rachlin, J.W. See Warkentine, B.E., 137
Price, W.W., 134 Radashevsky, V.I., 332
Pridmore, R. See Boyd, P.W., 456 Raffaelli, D. See Speirs, D., 136
Prince, R.I.T. See Walker, D.I., 475 Raffaelli, D.G. See Lawrie, S.M., 132
Probyn, T.A. See Attwood, C.G., 74 Rahbek, C. See Colwell, R.K., 167
Proctor, C. See Thresher, R., 474 Raiker, V. See Ramaswamy, V., 193
Proctor, T.D., 35 Railsback, S.F. See Grimm, V., 129
Prospero, J.M. See Jickells, T.D., 465 Raimondi, P.T., 84
Prostakova, I.V. See Shepelevich, N.V., 36 See Gaylord, B., 79
Provini, A. See Marchetti, R., 401 See Reed, D.C., 85
Prowse, T.A.A., 471 Rajan, P.T. See Turner, J.R., 194
Pruett, L., 403 Rajendran, N. See Kathiresan, K., 465
Prummel, W. See Lotze, H.K., 401 Ralph, P.J., 471
Przeslawski, R., 471 Ralston, S.V. See Wing, S.R., 88
Ptrachett, M.S. See Baird, A.H., 190 Ramaswamy, V., 193
Pubellier, M., 193 Ramírez, A., 333
Pueyo, S., 170 Ramírez, J.M. See Jerardino, A., 322
Pukaric, S.B.G.W., 403 Ramírez, M. See Correa, J.A., 312
Pulgar, J.M., 332 Ramon, M. See Pagès, F., 330
Pullin, A.S. See Sutherland, W.J., 338 Ramorino, L., 333
Punt, A.E. See Fulton, E.A., 461 Ramorino, L.M., 333
Purca, S. See Montecinos, A., 328 Ramos, A.A. See Badalamenti, F., 394
Purvis, A., 170 Ramos, A.M. See Letelier, S., 324
See Mace, G.M., 169 Ramos, M., 333
Puskaric, S. See Barmawidjaja, D., 394 See Ulloa, O., 340
519
AUTHOR INDEX
Ramos-Esplá, A.A. See Barbera, C., 394 Rex, M.A., 170

Ramsing, N.B. See Fossing, H., 316 See Levin, L.A., 169, 324
Rangin, C. See Pubellier, M., 193 Rey, F. See Erga, S.R., 460
Rao, K.H. See Ramaswamy, V., 193 Reyes, A. See Moreno, C.A., 328
Rao, N.S. See Ramaswamy, V., 193 Reyes, E. See Buschmann, A.H., 75, 308
Rao, P.S. See Ramaswamy, V., 193 Reyes, H. See Morales, C.E., 328
Rao, T.S.S., 193 Reyes, J. See van Waerebeek, K., 341
Raper, S. See Cubasch, U., 458 Reynolds, J.D. See Perry, A.L., 470
Rasheed, M. See Wild, C., 343 See Dulvy, N.K., 459
Rask, N., 403 Reyss, J.L. See Muñoz, P., 329
Raskin, R.G. See Costanza, R., 396, 457 Rhoads, D.C., 471
Rasmussen, J.B. See Vadeboncoeur, Y., 137 Ribbe, J. See Cai, W., 456
Rasmussen, K.L. See Petersen, K.S., 403 Ribera, C. See Delgado, L., 127
Ratilal, P. See Makris, N.C., 132 Ribera Maycas, E. See Macquart-Moulin, C., 132
Rautenberger, R. See Bischof, K., 307 Ribes, M., 134
Raven, J., 471 Rice, A.L., 135
Raven, J.A. See Beardall, J., 454 Rice, C.P. See Evans, M.S., 128
Ravilious, C. See Spalding, M.D., 194 Rich, H.W., 471
Ravussin, P.A. See Both, C., 455 Richards, D.V. See Davis, G., 458
Recasens, L. See Bozzano, A., 126 Richardson, A.J. See Bonnet, D., 455
Reddiah, K., 193 See Edwards, M., 459
Redlow, A.E. See Foley, A.M., 460 See Edwards, M.J., 459
Reed, C. See Edgar, G.J., 459 See Hays, G.C., 463
Reed, D.C., 84, 85, 333, 471
See Hobday, A.J., 464
See Amsler, C.D., 74
See Kirby, R.R., 466
See Eardley, D.D., 78
See McKinnon, A.D., 468
See Gaylord, B., 79
See Poloczanska, E.S., 407–478, 471
See Harrold, C., 80
Richardson, J.B. See Keefer, D.K., 322
See Hempel, W.M., 80
See Sandweiss, D.H., 334
See Raimondi, P.T., 84
Richardson, J.I. See Mrosovsky, N., 469
Reed, P.C. See Limpus, C.J., 467
Richardson, T.H. See Mrosovsky, N., 469
Rees, E.L.S. See Mackie, A.S.Y., 169
Richer de Forges, B., 471
Rehfisch, M.M. See Robinson, R.A., 472
Richoux, N.B., 135
Reid, C. See Aguilar, A., 305
Rick, I.P. See Modarressie, R., 469
Reid, D.D. See Lyle, J.M., 468
Rick, T.C. See Erlandson, J.M., 78
Reid, D.G. See Beare, D.J., 454
Reid, P.C. See Beaugrand, G., 454 Ricklefs, R.E., 171
See Frid, C., 398 Ridgway, K.R., 472
See Kirby, R.R., 466 Ridgwell, A.J. See Jickells, T.D., 465
Reid, R.J. See MacLeod, C.D., 468 Riebesell, U., 472
Reimers, C.E. See Komar, P.D., 35 See Engel, A., 460
Reiners, F. See Greve, W., 462 See Raven, J., 471
Reinsel, K.A. See Forward, R.B., 316 Riera, F. See Delgado, O., 396
Reipschläger A. See Pörtner, H.O., 471 Riera, T. See Carola, M., 126
Reise, K., 403 See Coma, R., 126
See Lotze, H.K., 401 Rigg, G.B., 85
Reisenbichler, K.R. See Kaufmann, R.S., 131 Riggio, S. See Badalamenti, F., 394
Reisewitz, S.E. See Beck, M.W., 395 Rijstenbil, J.W., 472
Relini, G. See Benedetti-Cecchi, L., 395 Rinaldi, A. See Simonini, R., 404
Remerie, T., 134 Rinde, E. See Christie, H., 396
Remis̆ová, V. See Menzel, A., 469 Rintoul, S. See Boyd, P.W., 456
Renaud, M.L. See Zein-Eldin, Z.P., 477 Rintoul, S.R. See Thresher, R., 474
Rendell, L., 333 Ripley, H.T. See Jaubert, J.M., 400
Renken, H. See Mumby, P.J., 469 Rippon, S., 403
Renshoff, S. See Fish, M.R., 460 Riquelme, V.A. See Buschmann, A.H., 308
Retamal, L. See Montecino, V., 327 Riseman, S.F. See Hutchins, D.A., 321
Retamal, M. See Gómez-Uchida, D., 317 Riser, K.L. See Dayton, P.K., 77, 396
Revenga, C. See Burke, L., 396 See Tegner, M.J., 87, 474
Revilla, E. See Grimm, V., 129 See Vilchis, L.I., 475
520
AUTHOR INDEX
Rismondo, A., 403 Roeckner, E. See Gregory, J.M., 462

Risoviç, D., 36 Roelfsema, C.M. See Duke, N.C., 459
Ritz, D.A., 135 Roemmich, D., 472
See Cohen, P.J., 126 See Willis, J.K., 476
See Flynn, A.J., 128 Roesler, C.S., 36
See Foster, E.G., 129 Roessig, J.M., 472
See Johnston, N.M., 130 Rogers, C.S., 472
See Metillo, E.B., 133 Rogers, K., 472
Rivadeneira, M.M, 333, 472 Rogers, P.J. See Ward, T.M., 476
See Moreno, R.A., 328 Rojas, O. See Loeb, V.J., 324
Rivalan, P., 472 Rojas, P.M., 333
Rivas, M. See Gutiérrez, D., 319 Roleda, M.Y. See Wiencke, C., 476
Rivera, G., 333 Rollins, H.B. See Sandweiss, D.H., 334
Rivera, P. See Westermeier, R., 343 Romaine, S. See Gattuso, J.P., 461
Rivera, P.J. See Westermeier, R., 343 Román, C. See Jaramillo, E., 322
Riveros, J.C. See Majluf., 325 Roman, D. See Correa, J.A., 312
Rizzo, M. See Barbera, C., 394 Romano, G. See Ianora, A., 321
Roa, R., 333 Romero, J. See Invers, O., 464
See Acuña, E., 304 See van Dalfsen, J. A., 405
See Gallardo, V.A., 316 Romero, L. See Tokeshi, M., 340
Roach, D.M. See Zaneveld, J.R.V., 36 Romme, W.H. See Norse, E.A., 170
Roast, S.D., 135 Romo, H. See Alveal, K., 305
Robbins, T.T. See Vilchis, L.I., 475 Ronan, C.E., 333
Roberts, C.M., 333
Ronnback, P. See Pihl, L., 403
See Balmford, A., 307
Roodbergen, S.P. See Both, C., 455
Roberts, D. See Kennedy, R.J.,
Rooney, N., 135
Roberts, D.A. See DeMartini, E.E., 77
Root, T.L., 472
See Hallacher, L.E., 80
Rosales, S.A. See Escribano, R., 315
See Plummer, K.M., 134
See Palma, W., 331
Roberts, J.D. See Bancroft, W.J.,
Rose, C. See Ene, A., 460
Roberts, J.M., 472
Rose, G.A., 472
Robertson, A.I. See Alongi, D.M., 453
Rosen, B. See Hughes, T.P., 464
Robertson, B.R. See Druehl, L.D., 78
Rosen, B.R., 193
Robertson, D.L. See Zimmerman, R.C., 88, 477
See Wilson, M.E.J., 194
Robertson, D.R. See Swearer, S.E., 339
Rosenbaum, K.L. See Norse, E.A., 170
Robertson, G. See Moreno, C.A., 328
Robinson, C. See Hays, G.C., 463 Rosenberg, M.S., 171
Robinson, R.A., 472 Rosenberg, R., 333, 403
See Learmonth, J.A., 466 See Baden, S., 394
Robinson, S. See Bonnet, D., 455 See Karlson, K., 400
Robison, B.H. See Kaufmann, R.S., 131 See Pihl L., 134
Robledo, D. See Hernández-Carmona, G., 80 Rosenblatt, R.H. See Stepien, C.A., 337
Robles, A. See Arntz, W.E., 306 Rosenbluth, B. See Rutllant, J.A., 334
Roca, E. See Sayre, R., 335 Rosenfield, A. See Mackenzie, C.L., Jr, 401
Rocamora, J. See Cano, J., 396 Rosenthal, R. See Dayton, P.K., 313
Rochelle-Newall, E. See Engel, A., 460 Rosenthal, R.J., 85
Rodbell, D.T. See Moy, C.M., 329 See Dayton, P.K., 76
Rodgers, K.B. See Orr, J.C., 470 Rosenthal, Y., 85
Rodolfo, K.S., 193 Rosenzweig, M.L., 171
Rodríguez, A. See Drake, P., 128 Rosenzweigk, C. See Root, T.L., 472
Rodríguez, L., 333 Ross, G.J.B., 472
See Escribano, R., 314 Ross, J. See Johnson, C., 465
See Marín, V.H., 326 Rosson, A. See Manríquez, P.H., 325
See Ortlieb, L., 330 Rost, B. See Riebesell, U., 472
Rodríguez, S.R., 85, 333 Rothlisberg, P., 472
Rodríguez-Graña, L., 333 Rothschild B. See Porteiro C., 471
Rodriguez-Montesinos, Y.E. See Hernández-Carmona, G., Rothstein, D. See Wilcove, D.S., 405
80, 81 Rotstayn, L.D. See Gordon, H.B., 462
Rodríguez-Prieto, C., 403 Rottiers, V. See Dewicke, A., 128
521
AUTHOR INDEX
Roughgarden, J., 85, 333 Rutten, L.M. See Fourqurean, J.W., 460
See Connolly, S.R., 311 Rvilious, C. See Spalding, M.D., 404
See Farrell, T.M., 315 Ryan, J. See Chavez, F.P., 311, 457
See Hughes, T.P., 464 Rylands, A.B. See Brooks, T.M., 395
Rouse, G.W. See Bertrand, Y., 166 Ryrholm, N. See Parmesan, C., 470
Roux, J.P. See Moloney, C.L., 327 Ryther, J.H., 334
Rovira, J., 334
Rowan, R., 193
Rowden, A. See Attrill, M.J., 166 S
Rowden, A.A. See Foggo, A., 167
Rowe, G.T. See Arntz, W.E., 306 Saalfeld, K. See Whitehead, P.J., 476
Rowley, R.J. See Ebeling, A.W., 78 Sabat, P., 334
See Harris, L.G., 80 Sabine, C.L. See Orr, J.C., 470
Rowling, K. See Thresher, R.E., 474 Sachdev, S. See Holyoak, M., 130
Sadovy, Y. See Dulvy, N.K., 459
Roy, C. See Cury, P., 312
Saenger, P. See Blasco, F., 455
Roy, D.B. See Hickling, R., 463
Safi, K. See Boyd, P.W., 456
Roy, K., 171, 334
Sagarin, R.D., 472
See Jablonski, D., 169
See Barry, J.P., 454
Rozbaczylo, N. See Castilla, J.C., 310
Sahli, J.M., 334
See Hernández, C.E., 319
Saigusa, M., 135
See Moreno, R.A., 328, 329
Saintilan, N., 472
See Quiroga, E., 332
Rogers, K., 472
Rubilar, P. See Moreno, C.A., 328
Saint-Jean, L. See Kouassi, E., 131
Rubolini, D. See Jonzén, N., 465
Saito, M., 334
Ruckelshaus, M. See Leslie, H., 324
Sakamoto, W. See Sato, K., 473
See Leslie, H., 400
Sala, E., 85, 334, 403
See Roberts, C.M., 333
See Kinlan, B.P., 82
Rud, N. See Petersen, K.S., 403
See Pandolfi, J.M., 402, 470
Rudi, E. See Baird, A.H., 190 See Kaufman L., 322
Rudolph, A. See Ahumada, R., 305 Salamanca, M. See Takesue, R.K., 339
Rudstam, L.G., 135 Salamanca, M.A. See Muñoz, P., 329
See Gal, G., 129 Salamanca, S.A. See Muñoz, P., 329
See Johannsson, O.E., 130 Salas, F., 171
See Smith, S.L., 136 Salazar, S. See Fariña, J.M., 315
Rue, E.L. See Hutchins, D.A., 321 Saldívar, M. See Takesue, R.K., 339
Ruedy, R. See Hansen, J., 462 Salemaa, H. See Kangas, P., 400
Ruiz, G.M. See Swearer, S.E., 339 Salen-Picard, C., 334
Ruiz, J. See Frere, E., 316 Saliba, L.J. See Jeftic, L., 400
Ruiz Fernandez, J.A. See Badalamenti, F., 394 Salinas, G.R. See Castro, L.R., 311
Ruiz-Fernandez, J.M. See Diaz-Almela, E., 458 Salinas, N., 85
Ruiz-Tagle, N., 334 Salinas, S. See Shaffer, G., 336
See Fernández, M., 316 See Strub, P.T., 338
Rumohr, H., 171 Salm, R.V. See Kaufman L., 322
Runge, J.A. See Kouwenberg, J.H.M., 466 Salo, H.M. See Markkula, S.E., 468
Rupp, G. See Uriarte, I., 340 Salzwedel, H. See Arntz, W., 306
Russell, F.S., 472 See Tarazona, J., 339
Russell, G.L. See Gregory, J.M., 462 Sameoto, D., 135
Russell, R., 171 Sameoto, D.D. See Longhurst, A.R., 132
See Menge, B.A., 327 Sammarco, P.W.,.473
Russo, G.F., 403 Samson, C.R. See Edgar, G.J., 459
Russo, R. See Bonaventura, R., 455 Samuel, M.D. See Harvell, C.D., 463
Ruth, M. See Berghahn, R., 395 San Vicente, C., 135
Rutllant, J. See Moraga, J., 328 Sánchez, M., 334
See Shaffer, G., 336 Sánchez, P. See Santelices, B., 335
See Strub, P.T., 338 Sanchez, R.P. See Porteiro C., 471
See Torres, R., 340 Sánchez Lizaso, J.L. See Badalamenti, F., 394
Rutllant, J.A., 334 Sánchez-Mata, A. See Barbera, C., 394
522
AUTHOR INDEX
Sanchez-Pinero, F., 334 Satyanarayan, C. See Turner, J.R., 194

Sanchez-Rodriguez, I. See Hernández-Carmona, G., 80, 81 Saunders, G.W., 85
Sanders, H., 334 See Druehl, L.D., 77, 78
Sanders, H.L., 171 See Lane, C.E., 82
See Allen, J.A., 166 Sautour, B. See David, V., 127
Sand-Jensen, K. See Marbà, N., 401 Sauvageau, C., 85
See Middelboe, A.L., 401 Sawamto, S., 135
Sandlin, P.E. See Bickmore, B.R., 33 Sawangarreruks, S. See Nielsen, T.G., 192
Sandulli, R., 403 Sawyer, T. See Mayer, L.M., 133
Sandweiss, D.H., 334 Sayre, R., 335
Sandweiss, M.D. See Sandweiss, D.H., 334 Scagel, R.F., 86
Sanger, G.A., 135 See Gabrielson, P.W., 79
Sangiorgi, F., 403 Scarabino, F. See Orensanz, J.M., 134
Sano, M. See Inoue, T., 130 Scarlett, B. See Heffels, C.M.G., 34
Sanpanich, K. See Turak, E., 194 Schaanning, M.T. See Berge, J.A., 455
Santander, H. See Carrasco, S., 309 Schaeffer, M.B. North, W.J., 84
Santelices, B., 85, 334, 335 Schaff, S. See Ene, A., 460
See Avila, M., 75, 306 Scheer, G., 193
See Bobadilla, M., 307 Scheffer, M. See Schippers, P., 473
See Castilla, J.C., 310 Scheibling, R.E., 473
See Correa, J.A., 312 Scheifinger, H. See Menzel, A., 469
See Hannach, G., 319 Schell, C. See De Robertis, A., 128
See Hoffmann, A., 320 Schell, P.A.,
See Hoffmann, A.J., 81, 320 Scheltema, R.S., 335
See Martínez, E.A., 326 Schembri, P.J. See Barbera, C., 394
See Meneses, I., 327 Scheridan, P.F. See Beck, M.W., 394
See Muñoz, M., 329 Scheuerell, M.D. See Schindler, D.E., 135
See Ojeda, F.P., 84, 330 Schiariti, A., 135
See Oliger, P., 330 Schick, L.L. See Mayer, L.M., 133
See Pizarro, A., 332 Schiecke, U. See Bacescu, M., 125
See Vásquez, J.A., 87, 341, 342 Schiel, D.R., 86, 473
See Villouta, E., 343 See Foster, M.S., 78, 79
Santibañez, A. See Gutierrez, A., 80, 319 Schiermeier, Q., 473
Santibáñez, J., 335 Schimmelmann, A., 86
Santibáñez, P. See González, H.E., 317 Schindler, D.E., 135
Santoro, C.M., 335 Schippers, P., 473
Santos, M.B. See Learmonth, J.A., 466 Schlacher, T.A., 135
Sanz, J.J. See Both, C., 455 Schlatter, R.P., 335
Sapper, J. See Zhao, T.X.-P., 36 Schlitzer, R. See Orr, J.C., 470
Sardiña, P., 135 Schmalfuss, H., 335
Sarmiento, J.L., 473 Schmid, M. See Kuypers, M.M., 323
See McNeil, B.I., 469 Schmidt, G.W. See LaJeunesse, T.C., 192
See Orr, J.C., 470 Schmidt, M. See Atkinson, L.P., 306
Sartor, P. See Bozzano, A., 126 Schmidt-Nielsen, K., 335
See Perez, T., 470 Schmiede, P. See Santelices, B., 335
Sartoretto, S. See Garrabou, J., 461 Schmitt, R.J. See Holbrook, S.J., 81, 464
Sarupia, J.S. See Desai, B.N., 191 Schmitz, K., 86
Satapoomin, S. See Nielsen, T.G., 192 Schneider, S.H. See Root, T.L., 472
Satapoomin, U., 193 Schneider, U. See Engel, A., 460
See Brown, B.E., 191 Schneider, W. See Atkinson, L.P., 306
See Chansang, H., 191 See Fuenzalida, R., 316
Sato, K., 473 See Palma, M., 331
Sato, M. See Hansen, J., 462 See Yannicelli, B., 344
Satta, M.P. See Martin, D., 401 Schoeman, D.S. See Richardson, A.J., 471
See Moschella, P.S., 402 Scholten, J. See Antia, A.N., 454
Satterlee, D.R. See Keefer, D.K., 322 Schonberg, S. See Dunton, K., 313
Satyabalam, S.P. See Bilham, R., 191 Schories, D., 403
523
AUTHOR INDEX
Schramm, W. See Vogt, H., 405 Setlow, J.K. See Setlow, R.B., 473
Schreurs, R. See Volten, H., 36 Setlow, R.B., 473
Schrieber, M.A. See Majluf, P., 325 Seva, A. See Barbera, C., 394
Schroeder, B.A. See Foley, A.M., 460 Sewell, J. See Hiscock, K., 399
Schroeter, S.C., 86 Sewell, R.B.S., 193
See Dean, T.A., 77 Seymour, J. See Little, M., 467
See Dixon, J., 77 Seymour, R.J., 86
See Reed, D.C., 85 Sfriso, A., 403, 404
Schul-Bull, D. See Antia, A.N., 454 Sgorbini, S. See Sandulli, R., 403
Schultz, D. See Norkko, A., 470 Shachak, M. See Jones, C.G., 322
Schulz, H. See Cowie, G.L., 312 Shaffer, G., 336
See Fossing, H., 316 See Hormazábal, S., 320
Schulz, H.N., 335 See Leth, O., 324
Schwartz, M.D., 473 Shah, D. See Madhaven, B.B., 192
See Wolfe, D.W., 477 Shanks, A.L., 86, 336
Schweder, C. See MacLeod, C.D., 468 See Mattison, J.E., 83
Schwindt, E. See Orensanz, J.M., 134 Shannon, C.E., 171
Scoffin, T.P., 193 Shannon, L.J. See Cury, P., 312
See Brown, B.E. 191 See Moloney, C.L., 327
See Tudhope, A.W., 194 Shaughnessy, G. See Ross, G.J.B., 472
Searle, M., 193 Shaughnessy, P.D. See Marsh, H., 468
Sebens, K.P., 335, 403, 473 See Ross, G.J.B., 472
Secretariat of the Convention on Biological Diversity., 335 Shaulis, N.J. See Wolfe, D.W., 477
Sedaghatkish, G. See Sayre, R., 335 Shchepetkin, A. See Marchesiello, P., 325
Seddon, P. See Ellenberg, U., 314 Shchepetkin, A.F., 336
Seddon, S., 473 Shears, N.T., 336
Sedwick, P.N., 473 Sheldon, R.W., 36
Seed, R. See Birkett, D.A., 395 Shelford, V.E., 171
Segrove, F. See Colman, J.S., 126 See Clements, F.E., 166
Seiderer, L.J. See Newell, R.C., 402 Shellito, S.M. See Abello, H.U., 124
Seikai, T. See Takeuchi, T., 136 See Taylor, L.H., 137
See Tominaga, O., 137 Shepelevich, N.V., 36
Seki, M.P. See Polovina, J.J., 471 Shepherd, J. See Raven, J., 471
Self, R.F.L. See Jumars, P.A., 131 Shepherd, S. See Johnson, C., 465
Selkoe, K.A. See Swearer, S.E., 339 See Sayre, R., 335
Sellanes, J., 335, 336 Sherman, R.K. See Lasenby, D.C., 132
See Gutiérrez, D., 319 Shi, G. See Cai, W., 456
See Milessi, A., 327 Shi, Y. See Lasenby, D.C., 132
See Muñoz, P., 329 Shick, J.M., 473
See Neira, C., 329 Shields, J.D., 135
See Thiel, M., 199–344 Shifrin, K.S., 36
Selmer, J.S. See Wängberg, S.A., 476 Shillington, F.A. See Hill, A.E., 320
Seltzer, G.O. See Moy, C.M., 329 Shima, J.S. See Swearer, S.E., 339
Semenov, V.N., 336 Shimode, S. See Kurihara, H., 466
Semroud, R. See Boudouresque, C.F., 395 Shirayama, Y. See Kurihara, H., 466
Senior, C.A. See Cubasch, U., 458 Short, F.T., 404, 473
Sepúlveda, F. See Simeone, A., 337 See Green, E.P., 398
Sepúlveda, H.H. See Thiel, M., 199–344 Shulenberger, E. See Haury, L.,
Sepulveda, J. See Pantoja, S., 331 Shumida, A. See Wilson, M., 172
Sepúlveda, M. See Sielfeld, W., 336 Shurin, J.B. See Leibold, M.A., 323
Sepúlveda, R., 336 Siebeck, O., 135
See Chong, J., 311 Siebeck, U.E., 473
SERNAPESCA, 336 Siegel, D.A., 36, 336
Serra, R. See Porteiro C., 471 Siegfried, C.A., 136
Serrão, E. See Brawley, S.H., 456 Sieh, K., 193
Serrão, E.A., 473 Sielfeld, W., 336, 337
Serviere-Zaragoza, E. See Hernández-Carmona, G., 80 Sievers, H. See Johnson, D.R., 322
Servos, M.R. See Johannsson, O.E., 130 Sifeddine, A. See Valdés, J., 340
Setchell, W.A., 86 Sigman, D.M. See Tortell, P.D., 475
524
AUTHOR INDEX
Sih, A., 404 Smith, G.J. See Coyer, J.A., 76

Silk, J.R. See Phillips, R.A., 332 See Hutchins, D.A., 321
Silva, C., 337 Smith, G.W. See Harvell, C.D., 463
See Yáñez, E., 344 Smith, I. See Allan, R., 190
Silva, M. See Palma, A.T., 331 Smith, J.E., 171
Silva, N., 337 Smith, J.R., 473
See Morales, C.E., 328 Smith, K.L., Jr. See Kaufmann, R.S., 131
Silva, P.C. See Moe, R.L., 83 Smith, L. See Babcock, R.C., 454
Silva-García, A. See Drake, P., 128 Smith, M.G. See McShane, P.E., 469
Silva-García, C. See Tognelli, M.F., 340 Smith, M.P.L. See Inglis, G.J., 464
Silverin, B. See Both, C., 455 Smith, R., 474
Silvestri, C. See Sandulli, R., 403 Smith, R.C. See Häder, D.P., 462
Simberloff, D.S., 171 Smith, R.D. See Griffies, S.M., 462
Simenstad, C. See Dean, A.F., 127 Smith, R.L. See Enfield, D.B., 314
Simenstad, C.A. See Duggins, D.O., 78 See Freeland, H.J., 316
Simeone, A., 337 See Huyer, A., 321
See Bernal, M., 307 Smith, S.D.A., 86
See Luna-Jorquera, G., 325 Smith, S.I., 136
See Schlatter, R.P., 335 Smith, S.L., 136
Simkanin, C., 473 Smith, W.O., Jr. See Barber, R.T., 125
Simonini, R., 404 Sneath, P.H.A., 171
Simpson, C.J., 473 Snelgrove, P.V.R., 171, 474
See Babcock, R.C., 454 See Quijón, P.A., 170
Snell, J.L. See Kemeny, J.G., 131
Simpson, E.H., 171
Snoeijs, P. See Choo, K.S., 457
Sims, D.W., 473
See Eriksson, B.K., 398
Singh, C.D. See Pal, T., 192
See Johansson, G., 400
Singham, S.B. See Bohren, C.F., 33
See Pedersén, M., 402
Sinke, J. See Dauwe, B., 127
Sobarzo, M., 337
Sinnassamy, J.M. See Boudouresque, C.F., 395
See González, H.E., 317, 318
Sinyuk, A. See Dubovik, O., 34
See Iriarte, J.L., 321
Siregar, A.M. See Baird, A.H., 190
See Pagès, F., 330
Sitts, R.M., 136
See Yannicelli, B., 344
Skeffington, M.J.S. See Curtis, T.G.F., 396
Sobel, J.A., 337
Skilleter, G.A. See Manson, F.J., 468
Sobolev, I., 337
Skillman, A.D. See Huesemann, M.H., 464
Sobrino, I. See Drake, P., 128
Skirving, W. See Done, T.J., 459 Soderqvist, T. See Pihl, L., 403
See Kleypas, J.A., 466 Soegiarto, A., 193, 194
Skirving, W.J. See Donner, S.D., 459 Soekarno, 194
Skottsberg, C., 86 Soimasuo, M.R. See Huovinen, P.J., 81
Slaper, H. See Kelfkens, G., 465 Sokal, R.R. See Sneath, P.H.A., 171
Slater, F.M. See Both, C., 455 Sokolov, L.V. See Both, C., 455
Slater, R. See Sarmiento, J.L., 473 Soldatov, V. See Sarmiento, J.L., 473
Slater, R.D. See Orr, J.C., 470 Solé, R.V. See Montoya, J.M., 133
Slavin, R.E., 171 Solow, A.R., 474
Slutsker, I. See Dubovik, O., 34 Solvang, R. See Jonzén, N., 465
Smart, J., 473 Somarakis, S. See Karakassis, I., 169
Smayda, T.J., 473 Somerfield, P.J., 171
Smedbol, R.K., 136 See Ellingsen, K.E., 167
Smetacek, V. See Ianora, A., 321 See Olsgard, F., 170
Smith, A.D.M. See Fulton, E.A., 461 See Warwick, R.M., 172
Smith, C.J. See Karakassis, I., 169 Somero, G.N., 337
Smith, C.R., 337 See Dahlhoff, E., 312
See Glover, A.G., 461 See Graves, J.E., 318
See Levin, L.A., 169, 324 See Hofmann, G.E., 320
See Snelgrove, P.V.R., 171, 474 See Stillman, J.H., 337
Smith, F. See Broitman, B.R., 308 Sommer, U. See Worm, B., 405
See Witman, J.D., 172 Sondergaard, M. See Aaser, H., 124
Smith, G.C. See Ross, G.J.B., 472 Song, K.-H., 136
525
AUTHOR INDEX
Sorbe, J.C., 136 Stanton, T.P. See Coale, K.H., 311

See Cartes, J.E., 126 Staples, D. See Rothlisberg, P., 472
See Cornet, M., 127 See Vance, D., 475
See Cunha, M.R., 127 Staples, D.J., 474
See Kouassi, E., 131 Stattegger, K. See Haneburth, T., 192
See Elizalde, M., 128 Stebbins, T.D., 86
Sotka, E.E., 337 Stech, J.L. See Zagaglia, C.R., 477
Soto, A. See Gutiérrez, D., 319 Steeman-Nielsen, E. See Jerlov, N.G.,
See Neira, C., 329 Steeves, T.E. See Darling, J.D., 127
Soto, R., 86, 337 Stefanescu, C. See Parmesan, C., 470
See Quiroga, E., 332 Stegenga, H., 86
Soto, V. See Martínez, G., 326 Steinbeck, J.R. See Schiel, D.R., 86
Soto-Gamboa, M. See Sabat, P., 334 See Schiel, D.R., 473
Sotomayor, A. See Navarrete, S.A., 329 Steinberg, P.D. See Estes, J.A., 78
See Wieters, E.A., 343 Steininger, F.F. See Harzhauser, M., 319
Southward, A. See Hiscock, K., 464 Stelle, L.L., 136
Southward, A.J., 474 Steller, D.L. See Zimmerman, R.C., 477
See Kendall, M.A., 465 Steneck, R.S., 86, 404, 474
See Mieszkowska, N., 469 See Adey, W.H., 453
See Simkanin, C., 473 See Jackson, J.B.C., 399
See Sims, D.W., 473 See Vadas, R.L., 475
Spalding, H., 86, 474 Stenseth, N.C. See Jonzén, N., 465
Spalding, M. See Burke, L., 396 Stephen, A.C., 171
Spalding, M.D., 194, 404
Stephens, J.S., Jr, 86
Spall, S.A. See Sarmiento, J.L., 473
See Holbrook, S.J., 464
Span, A. See Boudouresque, C.F., 395
Stephenson, R. See Smedbol, R.K., 136
Spanhoff, R. See Hamm, L., 399
Stephensons, C. See Conkright, M.E., 311
Spärck, R., 171
Stepien, C.A., 337
Sparks, T.H. See Lehikoinen, E., 467
Stergiou, K.I. See Browman, H.I., 395
See Menzel, A., 469
Stervander, M. See Jonzén, N., 465
See Robinson, R.A., 472
Stevens, C.L. See Hurd, C.L., 81
Sparrow, A.D. See Friedel, M.H., 461
Stewart, H.L., 474
Speirs, D., 136
Stewart, J.E. See Holmer, M., 399
Speirs, D.C. See Lawrie, S.M., 132
Steyaert, M. See Deprez, T., 128
Spencer, C.N., 136
St-Hilaire, A., 136
Spencer, M. See Kiflawi, M., 169
Spencer, T., 194 Stillman, J.H., 337
Speransky, V. See Brawley, S.H., 456 Stimson, J. See Stambler, N., 194
Speybroeck, J. See Deprez, T., 128 Stive, M.J.F. See Hamm, L., 399
Spicer, J.L., 337 Stoddart, D.R. See Ellison, J.C.,460
Spiers, A.G. See Finlayson, C.M., 398 Stokes, T. See Ross, G.J.B., 472
Spiess, F.N. See Genin, A., 461 Stone, C.J., 337
Spina, F. See Jonzén, N., 465 Stone, H.H., 136
Spinrad, R.W. See Codispoti, L.A., 311 Stone, R. See Allan, R., 190
Spivak, E.D. See Orensanz, J.M., 134 Stoner, D. See Roughgarden, J., 333
Spotila, J.R. See Binckley, C.A., 455 Stoner, D.S., 474
Srinivasan, A. See Cowen, R.K., 457 Stott, P.A. See Karoly, D.J., 465
Stachowicz, J.J. See Byrnes, J., 75 Stotz, L.M. See Caillaux, L.M. 309
Stafford, R. See Attrill, M.J., 166 Stotz, W., 337, 338
Stambler, N., 194 See Aburto, J., 304
Stamnes, J.J, See Erga, S.R., 460 See González, J., 318
Stamnes, K. See Erga, S.R., 460 See González, S.A., 318
Standen, V.G. See Briones, L., 308 See Jaramillo, E., 322
See Santoro, C.M., 335 See Jesse, S., 322
Standford, J.A. See Spencer, C.N., 136 See León, R.I., 324
Stanford, J.A. See Chess, D.W., 126 See Meltzoff, S.K., 327
Stanley, G.D., 194 See Ortiz, M., 330
Stanners, D., 404 See Pérez, E.P., 331
Stanton, T.K. See Lawson, G.L., 132 See Thiel, M., 199–344
526
AUTHOR INDEX
See von Brand, E., 343 Sutherland, W.J., 338

See Zamora, S., 344 Sutton, C.W. See Eardley, D.D., 78
Stotz, W.B. See Aguilar, M., 305 See Hempel, W.M., 80
See Caillaux, L.M., 309 Sutton, P. See Boyd, P.W., 456
See Lancellotti, D.A., 323 See Costanza, R., 396, 457
See Lopez, C.A., 324 Svendsen, E. See Edwards, M.J., 459
See Pacheco, A., 330 Swadling, K.M. See Foster, E.G., 129
Stouffer, R. See Sarmiento, J.L., 473 Swanson, A.K., 86, 474
Stouffer, R.J. See Cubasch, U., 458 Swearer, S.E., 339
See Gregory, J.M., 462 Sweeney, C. See Langdon, C., 466
Stout, M.L. See Norse, E.A., 170 Swenarton, J.T. See Keser, M., 466
Stow, D.A. See Augenstein, E.W., 74 Sykes, M.T. See Walther, G.R., 476
St-Pierre, J.F. See Kouwenberg, J.H.M., 466 Sylvester, C. See Grünewald A., 318
Stramski, D., 36 Symonds, D.T. See Makris, N.C., 132
See Boss, E., 33 Syvitski, J.P.M., 36
Strathmann, R.R., 338
See Cohen, C.S., 311
See Lee, C.E., 323
See Palmer, A.R., 331
T
Striz, M. See Menzel, A., 469 Taghon, G.L. See Mayer, L.M., 133
Strong, A.E. See Kleypas, J.A., 466 Taguchi, S. See Goes, J.I., 462
Strotmann, B. See Schulz, H.N., 335 Takahashi, K., 136
Strub, P.T., 338 Takahashi, T. See Langdon, C., 466
See Blanco, J.L., 307 Takesue, R.K., 339
See Carr, M.E., 309 Takeuchi, T., 136
See Halpin, P.M., 319 Tala, F., 339
See Hill, A.E., 320 See Edding, M., 314
See Mesías, J.M., 327 See Thiel, M., 199–344
See Montecino, V., 327 See Vásquez, J.A., 342
See Thomas, A.C., 339 See Véliz, K., 342
Strzepek, R. See Boyd, P.W., 456
Tammaru, T. See Parmesan, C., 470
Stuardo, J., 338
Tan, I.H. See Druehl, L.D., 78
Stuart, C.T. See Gage, J.,167
Tanaka, H. See Sato, K., 473
See Levin, L.A., 169, 324
Tanaka, M. See Islam, M.S., 399
See Rex, M.A., 170
See Tanaka, Y., 136
Stuck, L. See Price, W.W., 134
Tanaka, Y., 136
Stuxburg, A., 171
Tanguyt, A. See Moragat, D., 328
Su, J.C., 171
Tankersley, R.A. See Forward, R.B., 316
Su, M. See Ene, A., 460
Tanneberger, K. See Boyd, P.W., 456
See Herbst, L., 463
Subba Rao, N.V. See Turner, J.R., 194 Tanner, S.J. See Martin, J.H., 326
SUBPESCA, 338 Tanré, D. See Zhao, T.X.-P., 36
Suchanek, T.H., 404 Tapia, C. See González, J., 318
Suda, Y. See Inoue, T., 130 Tapia, F. See Lagos, N.A., 323
Suh, H.-L., 136 See Roa, R., 333
Suharsono, 194 Tararam, A.S., 137
Suhr, S.B. See Gooday, A.J., 318 Tarazona, J., 339
Summerville, K.S. See Veech, J.A., 172 See Arntz, W., 306
Sundareshwar, P.V. See Morris, J.T., 402 See Fernández, E., 315
Sundelof, A. See Moschella, P.S., 402 See Montecino, V., 327
See Airoldi, L., 394 See Peña T.S., 331
Surman, C.A., 474 See Rosenberg, R., 333
See Dunlop, J.N., 459 Targett, T.E. See Lankford, T.E., 132
Suryanarayana, A. See Varkey, M.J., 194 Tarifeño, E. See Castro, L.R., 311
Susnik, A. See Menzel, A., 469 See Chong, J., 311
Sutcliffe, W.H., Jr. See Sheldon, R.W., 36 Tarling, G.A. See Emsley, S.M., 128
Sutherland, D.L., 136 Tasker, M. See Frid, C., 398
Sutherland, J.P. See Moreno, C.A., 328 Tattersall, W.M., 137
See Moreno C.A., 83 Taylor, J.H. See Hunter, J.R., 464
527
AUTHOR INDEX
Taylor, L.H., 137 Thompson, R.J. See MacDonald, B.A., 325

See Abello, H.U., 124 See Richoux, N.B., 135
Teare, J.A. See Simeone, A., 337 Thomsen, H.A. See Nielsen, T.G., 192
Teas, W.G. See Foley, A.M., 460 Thomson, B.E. See Worrest, R.C., 477
Tegen, I. See Jickells, T.D., 465 Thomson, D.L. See Crick, H.Q.P., 457
Tegner, M.J., 86, 87, 474 Thoney, D.A. See Drohan, A.F., 459
See Dayton, P.K., 76, 77, 396, 458 Thornber, C.S. See Byrnes, J., 75
See Jackson, J.B.C., 399 Thorne, R.E., 137
See Schimmelmann, A., 86 Thorpe, A. See Aguilar, A., 305
See Seymour, R.J., 86 Thorrold, S.R. See McKinnon, A.D., 468
See Steneck, R.S., 86, 404, 474 See Swearer, S.E., 339
See Vilchis, L.I., 475 Thorson, G. 171
Tennent, W.J. See Parmesan, C., 470 Thorson, G.L., 339
Terborgh, J.W., 171 Thresher, R., 474
Terbrueggen, A. See Engel, A., 460 Thresher, R.E., 474
Terlizzi, A. See Guidetti, P., 398 Thrush, S.F., 171, 404, 474, 475
Terrazzano, G. See Ianora, A., 321 See Lohrer, A.M., 467
Terreay, L. See Bopp, L., 455 See Norkko, A., 470
Teske, A. See Fossing, H., 316 See Turner, S.J., 404
Teske, P.R., 339 Thulke, H.-H. See Grimm, V., 129
Tester, P. See Carlsson, P., 456 Thurston, M.H., 340
Tett, P.B. See Gage, J., 167 Thwin, S. See Ramaswamy, V., 193
Thamdrup, B. See Fossing, H., 316 Tija, H.D., 194
Thangaduri, G. See Kathiresan, K., 465 Tilman, D. See Leibold, M.A., 323
Theroux, R.B. See Wigley, R.L., 138 Tinney, L. See Jensen, J.R., 81
Thessen, A.E., 474 Tintore, J. See Marbà, N., 401
Thibaut, T., 404 Tiselius, P. See Peterson, W.T., 331
See Meinesz, A., 401 Tittley, I. See Bartsch, I., 394
Thiel, M., 87, 199–344, 339 See Hiscock, K., 464
See Hinojosa, I., 320 Tognelli, M.F., 340
See Macaya, E.C., 83, 325 Tokeshi, M., 340
See Sepúlveda, R., 336 tom Dieck, I. See Lüning, K., 325
See Valdivia, N., 340 See Wiencke, C., 343
See Wahl, M., 475 Tomanek, L., 340
Thieltges, D.W. See Reise, K., 403 Tomas, F. See Invers, O., 464
Thies, K. See Dean, T.A., 77 Tomasin, A. See Bondesan, M., 395
See Schroeter, S.C., 86 Tomczak, M., 194
Thivierge, D. See Winkler, G., 138 Tomicic, J.J., 340
Thomas, A. See Thiel, M., 199–344 Tominaga, O., 137
Thomas, A.C., 339 See Tanaka, Y., 136
See Blanco, J.L., 307 See Yamamoto, M., 138
See Carr, M.E., 309 Tompkins, D.M. See Mouritsen, K.N., 469
See Hill, A.E., 320 Torbjørn, E. See Jonzén, N., 465
See Montecino, V., 327 Torok, J. See Both, C., 455
Thomas, C.D., 474 Torras, X. See Thibaut, T., 404
See Hickling, R., 463 Torres, R., 340
See Parmesan, C., 470 See Jickells, T.D., 465
Thomas, D. See Harris, G., 463 See Rutllant, J.A., 334
Thomas, J.A. See Parmesan, C., 470 Torres-Villegas, J.R. See Hernández-Carmona, G., 80, 81
Thomas, L.P., 474 Tortell, P.D., 475
Thomason, J.C. See Wahl, M., 475 See Riebesell, U., 472
Thompson, D.W. See Gillett, N.P., 461 Totterdell, I.J. See Orr, J.C., 470
Thompson, J.B. See Mullins, H.T., 329 Tourre, Y. See Allan, R., 190
Thompson, R.C., 404 Tourre, Y.M. See White, W.B., 476
See Airoldi, L., 394 Tovar, H., 340
See Jonsson, P.R., 465 Towle, D.W., 87
See Martin, D., 401 Tranter, J. See Ross, G.J.B., 472
See Mieszkowska, N., 469 Travers, V. See Cole, R.G., 457
See Moschella, P.S., 402 Travis, J.M.J., 475
528
AUTHOR INDEX
Travis, L.D. See Mishchenko, M.I., 35 UK Biodiversity Group, 404

Treleani, R. See Caressa, S., 396 UK Steering Group, 404
Trenberth, K.E., 340 Ulloa, N. See Gómez, I., 317
Trent, J.D. See Mattison, J.E., 83 See Huovinen, P., 321
Trick, C.G. See Hutchins, D.A., 321 Ulloa, O., 340
Trillmich, F., 340 See Castilla, J.C., 310
See Dellinger, T., 313 See Daneri, G., 312
Trinder, M. See Votier, S.C., 475 See Escribano, R., 314
Troell, M. See Chopin, T., 311 See Farías, L., 315
See Pihl, L., 403 See Fossing, H., 316
Trombini, C., 404 See Hormazábal, S., 320
Troncoso, A. See Bernal, P.A., 307 See Leth, O., 324
See González, H.E., 317 See Yuras, G., 344
Troncoso, L. See Galleguillos, R., 316 Ullrich, N. See Thiel, M., 339
Troncoso, V.A., 340 Umani, S.F. See Bonnet, D., 455
See Castro, L.R., 310 UNEP/FAO/WHO, 404
See Iriarte, J.L., 321 UNEP/IUCN, 194
Trull, T. See Boyd, P.W., 456 UNEP/MAP/PAP, 404
Trull, T.W. See Sedwick, P.N., 473 UNEP/WCMC, 405
Trumbore, S. See Mayer, L.M., 133 United Nations Conference on Environment and
Tsuchiya, M., 475 Development, 172
Tsuei, T.G. See Geller, P.E., 34 Uno, N. See Tominaga, O., 137
Tsukayama, I. See Pauly, D., 331 Urban, H.J., 340
Tsusaki, T. See Tanaka, Y., 136 Uriarte, I., 340
Tudela, S., 137, 404 Uribe, E., 340Uriarte A. See Porteiro C., 471
Tudhope, A.W., 194 See Acuña, E., 305
See Brown, B.E., 191 See Avaria S., 306
See Scoffin, T.P., 193 See Illanes, J.E., 321
Tunesi, L. See Meinesz, A., 401 Uribe, M. See Castilla, J.C., 310
Turak, E., 194 Urzua, C. See Vásquez, J.A., 342
See De Vantier, L.M., 458 Utter, B.D., 87
Turley, C. See Raven, J., 471
Turner, C.H. See Feder, H.M., 78
Turner, D.R. See Torres, R., 340 V
Turner, I.M. See Hesp, P.A., 192
Turner, J.R., 194 Vacelet, J. See Perez, T., 470
Turner, S. See Boyd, P.W., 456 Vadas, R.L., 475
Turner, S.J., 404 Vadeboncoeur, Y., 137
Turpen, S.L. See Anderson, B.S., 74 Väinölä, R. See Audzijonyte, A., 125
Tutschulte, T.C., 87 Valdebenito, E. See Moraga, J., 328
Twardowski, M.S., 36 Valdés, J., 340
See Boss, E., 33 Valdes, J. See Ortlieb, L., 330
Twilley, R.R., 475 Valdes, L. See Bonnet, D., 455
See Ewel, K.C., 460 Valdivia, E. See Arntz, W.E., 306
Twining, B.S., 137 Valdivia, N., 340
Tyagi, A.P., 475 See Hinojosa, I., 320
Tyler, P.A., 172 Valdivia, N.A. See Macaya, E.C., 325
Tyler-Walters, H., 404 Valdivia, N.E. See Macaya, E.C., 83
See Hiscock, K., 399 Valdovinos, C., 172, 340
Tyson, R.V., 340 See Castilla, J.C., 310
See Stuardo, J., 338
U Valdovinos, C.R. See Fernández, M., 315
Valentine, J.F. See Mattila, J., 132
Udy, J.W. See Abal, E.G., 453 Valentine, J.P., 475
Ugalde, P. See Hinojosa, I., 320 Valentine, J.W. See Jablonski, D., 169
Ugarte, R. See Hoffmann, A.J., 320 Valentine, W.V. See Roy, K., 171
Ugland, K. I., 172 Valenzuela, G. See Vargas, C.A., 341
See Gray, J.S., 168 Valenzuela, G.S. See Vargas, C.A., 341
529
AUTHOR INDEX
Valenzuela, J. See Alveal, K., 305 Varas, M. See Castilla, J.C., 310
Valenzuela, V. See Balbontín, F., 306 See Correa, J.A., 312
Valero, J. See González, J., 318 See Takesue, R.K., 339
Valero, M. See Faugeron, S., 315 Varela, C. See Jaramillo, E., 322
Valiela, I., 405 Varela, D. See Buschmann, A.H., 308
See Hauxwell, J., 399 Vargas, C.A., 341
Vallarino, E.A. See Orensanz, J.M., 134 See González, H.E., 317
Valle, C. See Barbera, C., 394 See Narváez, D.A., 329
Vallejo, L. See Marín, V.H., 326 See Piñones, A., 332
Valle-Levinson, A., 341 See Thiel, M., 199–344
See Atkinson, L.P., 306 Vargas, M. See Sielfeld, W., 336, 337
See Piñones, A., 332 Vargo, G.A. See Okey, T.A., 470
See Yannicelli, B., 344 Varkey, M.J., 194
Vallet, C., 137 Vasconcellos, M. See Okey, T.A., 470
See Dauvin, J.-C., 127 Vásquez, C. See Gutiérrez, D., 319
See Zouhiri, S., 138 Vasquez, J.A. See Thiel, M., 87, 199–344, 339, 341, 342
van Aken, H.M. See Philippart, C.J.M., 470 See Buschmann, A.H., 75, 308
Van Beusekom, J. See Lotze, H.K., 401 See Camus, P.A., 309
van Beusekom, J.E.E., 405 See Fernández, M., 315
van Bressem, M. See van Waerebeek, K., 341 See González, J., 318
Vance, D., 475 See Graham, M.H., 39–88, 318
Vance, D.J., 475 See Lancellotti, D.A., 323
See Staples, D.J., 474 See Macaya, E.C., 83, 325
Van Cuyck, J.P. See David, P.M., 127 See Munoz, V., 83, 329
van Dalfsen, J. A., 405 See Nuñez, L., 84, 330
van de Hulst, H.C., 36 See Santelices, B., 335
van den Belt, M. See Costanza, R., 396, 457 See Tala, F., 339
van den Berg, J.B., 405 See Thiel, M., 87
Vanden Berghe, E. See Deprez, T., 128 See Vega, J.M.A., 88, 342
van den Hoff, J. See Lake, S., 132 Vásquez, N. See Hinojosa, I., 320
van der Baan, S.M., 137 Vásquez, N.R. See Macaya, E.C., 83, 325
Vanderhoe, H. See Harris, G.P., 463 Vassen, W. See Volten, H., 36
van der Leun, J.C. See Kelfkens, G., 465 Vassura, I. See Trombini, C., 404
van der Velde, G. See Dorenbosch, M., 459 Vasta, G.R. See Harvell, C.D., 463
van der Zande, W.J. See Hoovenier, J.W., 34 Veas, R. See Palma, A.T., 331
Vander Zanden, M.J. See Vadeboncoeur, Y., 137 Veech, J.A., 172
van der Zwaan, G. See Barmawidjaja, D., 394 See Crist, T.O., 167
van Dijken, G.L. See El-Sayed, S., 459 Vega, A. See Buschmann, A.H., 75
van Donk, E. See Hessen, D., 463 See Buschmann, A.H., 308
Van Dover, C.L. See Doerries, M.B., 167 See Shaffer, G., 336
Van Dyke, H. See Karanas, J.J., 465 Vega, C. See Medina, M., 326
See Worrest, R.C., 477 Vega, J.M.A., 88, 342
Vanfleteren, J. See Remerie, T., 134 See Macaya, E.C., 83, 325
van Geen, A. See Takesue, R.K., 339 See Vásquez, J.A., 87, 341, 342
van Loveren, H. See Kelfkens, G., 465 Vega, M.A. See Letelier, S., 324
van Oijen, T. See Kelfkens, G., 465 See Vásquez, J.A., 341
Vanreusel, A. See Remerie, T., 134 Vega, S.A. See Rutllant, J.A., 334
Van Rijswijk, P. See Dauwe, B., 127 Veit, R.R. See Helmuth, B.S., 80
Van Tresca, D. See Dayton, P.K., 76 Velasco-Hernández, J.X. See Velázquez, I., 342
Van Tüssenbroek, B.I., 87 Velázquez, I., 342
See Cruz-Palacios, V., 457 Velders, G.J.M. See Kelfkens, G., 465
van Vierssen, J. See Kabat, P.W., 400 Véliz, D., 342
van Vliet, A.J.H. See Menzel, A., 469 See Vásquez, J.A., 87, 342
van Waerebeek, K., 341 Véliz, F. See Lagos, N., 323
See Guerra, C., 318 See Wieters, E.A., 343
van Woesik, R. See LaJeunesse, T.C., 192 Véliz, K., 342
Váradi, L., 405 Vellanoweth, R.L. See Erlandson, J.M., 78
530
AUTHOR INDEX
Vellinga, P. See Kabat, P.W., 400 Viviani, C.A., 88, 343

Veloso, X. See Sielfeld, W., 336 Vogt, H., 405
Ven Tresca, D. See Dayton, P.K., 313 Voicu, G., 137
Venegas, M. See Edding, M., 313, 314 Volhardt, C. See Negri, A., 469
Venegas, R.M. See Navarrete, S.A., 329 Vollmer, M. See Lotze, H.K., 401
See Vargas, C.A., 341 Volse, L.A. See Devinny, J.S., 77
Venkataraman, G. See Madhaven, B.B., 192 Volten, H., 36
Venkataraman, K. See Turner, J.R., 194 See Hoovenier, J.W., 34
Venrick, E.L., 475 von Brand, E., 343
Veraart, J. See Kabat, P.W., 400 von Oertzen, J.A. See Messner, U., 401
Vercoutere, T.L. See Mullins, H.T., 329 von Rad, U. See Cowie, G.L., 312
Vergara, K. See Schmalfuss, H., 335 Voss, K.J., 36
Vergara, P.A. See Buschmann, A.H., 308 See Stramski, D., 36
Vergara, S. See Correa, J.A., 312 Votier, S.C., 475
Verheijen, P.J.T. See Heffels, C.M.G., 34 Voultsiadou, E. See Badalamenti, F., 394
Verheye, H.M., 342 Vousden, D. See Turner, J.R., 194
See Cury, P., 312 Vuorinen, I. See Flinkman, J., 128
See Richardson, A.J., 471
Verlaque, M. See Boudouresque, C.F., 395
See de Villèle, X., 397 W
Vernberg, F.J., 342
Wabnitz, C.C.C. See Mumby, P.J., 469
Veron, C. See Turak, E., 194
Wada, T. See Porteiro C., 471
Vetter, E.W., 88, 342
Wærn, M. See Kautsky, N., 400
Vidal, J., 343
Wagener, A. See Mock, G., 402
Vidal, M.A. See Moreno, R.A., 328
Wahl, M., 475
Vierstraete, A. See Remerie, T., 134
See Valdivia, N., 340
Viherluoto, M., 137
Wahle, R.A., 137
See Viitasalo, M., 137
Waite, A. See Boyd, P.W., 456
Viitasalo, M., 137 Wakabara, Y. See Tararam, A.S., 137
See Linden, E., 132 Waldenström, J. See Jonzén, N., 465
See Viherluoto, M., 137 Walker, D., 475
Vik, J.O. See Jonzén, N., 465 Walker, D.I., 475
Vilchis, L.I., 475 Walker, T.A., 475
Vilchis, M.A. See Hernández-Carmona, G., 80, 81 Walla, T.R., Lande, R., 169
Vilina, Y. See Capella, J., 309 Wallace, C.C., 194
Vilina, Y.A. See Frere, E., 316 See Babcock, R.C., 454
Villablanca, R. See Simeone, A., 337 Wallace, J.M. See Mantua, N.J., 468
Villafane, V.E. See Barbieri, E.S., 454 See Zhang, Y., 344
Villagrán, C., 343 Wallace, R.S. See Simeone, A., 337
See Hinojosa, L.F., 320 Wallem, K.P. See Fariña, J.M., 315
See Maldonado, A., 325 Walsh, A. See Bowman, D.M.J.S., 456
See VillaMartínez, R., 343 Walsh, J.J., 343
VillaMartínez, R., 343 Walsh, K., 475
Villanueva, R. See Navarro, J.C., 133 See Pittock, A.B., 470
Villarroel, J.C., 343 Walsh, K.J.E., 476
See Acuña, E., 305 Walters, C.J. See Hilborn, R., 320
Villarroel, P. See Pauchard, A., 331 Walther, G.R., 476
Villegas, M., 343 Wang, H.J., 343
Villegas, M.J., 88 Wang, S.W. See Gong, D.Y., 317
Villouta, E., 343 Wang, Z., 137
Vincent, W.F. See Winkler, G., 138 See Vallet, C., 137
Vincx, M. See Deprez, T., 128 Wang Chen, H. See Alpers, W., 190
See Dewicke, A., 128 Wängberg, S.A., 476
See Hampel, H., 129 Ward, B.B. See Codispoti, L.A., 311
Vinogradov, M.E., 137 Ward, D. See MacNally, R., 169
Visser, A.W., 137 Ward, J.R., 476
Visser, M.E. See Both, C., 455 See Harvell, C.D., 463
531
AUTHOR INDEX
Ward, T.M., 476 Weissner, R. See Vásquez, J.A., 342

See Marsh, H., 468 Wellington, G.M., 476
Ware, D.M., 476 Wells, F.W., 476
Waren, A. See Rex, M.A., 170 Wells, S. See Kelleher, G., 400
Wares, J.P., 343 Welsford, D.C., 476
Warkentine, B.E., 137 Wennhage, H. See Pihl, L., 403
Warner, R.R. See Airamè, S., 393 Wenzel, H. See Westermeier, R., 343
See Jackson, J.B.C., 399 Wernberg, T. See Fowler-Walker, M.J., 460
See Pandolfi, J.M., 402 West, G. See Polacheck, T., 471
See Roberts, C.M., 333 West, L.D. See Lyle, J.M., 468
See Swearer, S.E., 339 West, R.J., 476
See Zacherl, D.C., 344 Westermeier, R., 88, 343
Warren, M. See Parmesan, C., 470 Westermeier, R.C. See Gómez, I., 317
Warwick, R.M., 172 Weston, M.A. See Chambers, L.E., 457
See Clarke, K.R., 166, 191 Westphal, C.J. See Clarke, P.J., 457
See Ellingsen, K.E., 167 Whang, A. See Janssen, J., 130
See Gobin, J.F., 168 Wheeler, A.J. See Roberts, J.M., 472
Washburn, L. See Gaylord, B., 79 Wheeler, P.A., 88
See Raimondi, P.T., 84 Wheeler, W.N., 88
See Reed, D.C., 85 See Druehl, L.D., 78
Watanabe, J.M., 88 Whetton, P. See Done, T.J., 459
See Graham, M.H., 79 Whitaker, A. See De Grave, S., 396
Waterman, L.J. See Gordon, H.B., 462 Whitakker, R.H., 172
Waterman, P.C., 36 White, E.P. See Hurlbert, A.H., 168
Waterman, T.H., 137 White, J.R. See Parkinson, R.W., 470
See Bainbridge, R., 125 White, K.N. See Crisp, D.J., 312
Waters, L.B.F.M. See Hoovenier, J.W., 34 White, R. See Mock, G., 402
Watkins J.L. See Brierley A.S., 126 White, W.B., 476
Watkinson, A.R. See Fish, M.R., 460 Whitehead, H. See Rendell, L., 333
See Gardner, T.A., 461 Whitehead, P.J., 476
Watling, L., 405 Whiteley, G.C., Jr, 138
See Thiel, M., 339 Whitfield, J., 172
Watson, A. See Raven, J., 471 Whitmarsh, D. See Badalamenti, F., 394
Watson, A.J. See Boyd, P.W., 456 Wichard, T. See Ianora, A., 321
Watson, E. See Gelcich, S., 317 Wickett, M.E. See Caldeira, K., 456
Watson, L.M. See Pomeroy, R.S., 332 Wicklum, D., 138
Watterson, I.G. See Gordon, H.B., 462 Widdicombe, S., 172
Way, L.S. See Davidson, N.C., 396 See Austen, M.C., 166
Waycott, M. See Carruthers, T.J.B., 457 Widdows, J. See Roast, S.D., 135
Wearmouth, V.J. See Sims, D.W., 473 Widdowson, T.B. See Gabrielson, P.W., 79
Weatherbee, R. See Thomas, A.C., 339 Widmann, M. See Jones, J.M., 465
Weaver, R.S. See Hutchins, D.A., 321 Wiebe, P.H. See Haury, L.R., 319
Webb, P., 137 See Lawson, G.L., 132
Weber, U. See Brandt, A., 126 Wiegand, T. See Grimm, V., 129
Webster, M.S. See Menge, B.A., 327 Wielgolaski, F-E. See Menzel, A., 469
Webster, P.J, 194, 476 Wiencke, C., 343, 476
Weeks, S.J., 343 See Bischof, K., 455
See Bakun, A., 306 See Hanelt, D., 462
Weetman, D. See Gómez-Uchida, D., 317 Wieters, E.A., 343
Wefer, G., 319 See Navarrete, S.A., 329
Weichler, T., 343 Wiff, R. See Quiroz, J.C., 332
Weidman, C. See Thresher, R., 474 Wigley, R.L., 138
Weikert, H. See Bonnet, D., 455 See Maurer, D., 133
Weiner, J. See Grimm, V., 129 Wilcove, D.S., 405
Weiner, N., 172 See Norse, E.A., 170
Weinstein, M.P. See Beck, M.W., 394 Wilcox, B.A. See Norse, E.A., 170
Weirig, M.F. See Orr, J.C., 470 Wild, C., 343
Weishampel, J.F., 476 Wildish, D.J. See Holmer, M., 399
Weissing, F.J. See Huisman, J., 464 Wildman, R.D. See Dawson, E.Y., 76
532
AUTHOR INDEX
Wilhelm, C., 476 Wolfe, D.W., 477

Wilhelm, F.M., 138 Wolff, M., 344
Wilkerson, F.P. See Hogue, V.E., 464 See Ortiz, M., 330
Wilkie, M.L., 405 Wolff, W.J., 405
Wilkin, J.L. See Griffin, D.A., 462 See Lotze, H.K., 401
Wilkinson, C., 194, 476 Wolters, M., 405
Wilkinson, C.R., 405, 476 Womersley, H.B.S., 88, 477
Williams, A., 476 Wones, A. See Behrenfeld, M., 455
See Koslow, J.A., 466 Wood, L.J., 344
Williams, A.B., 138 Wood, S.A. See Russell, R., 171
Williams, C.B. See Fisher, A.A., 167 Woodle, C.M. See Roessig, J.M., 472
Williams, K. See Majkowski, J., 468 Woodley, C.M. See Downs, C.A., 191
Williams, L. See Manson, F.J., 468 Woodroffe, C., 477
Williams, P.H., 172 Woodworth, P.L. See Church J.A., 457
See Gaston, K.J., 168 Wooldridge, S. See Done, T.J., 459
Williams, R. See Green, K., 129 Wooldridge, T.H. See Jerling, H.L., 130
Williams, R.J. See Dunne, J.A., 128 See Schlacher, T.A., 135
See Evans, M.J., 460 See Webb, P., 137
See Saintilan, N., 472 Wooller, R.D. See Dunlop, J.N., 459
Williams, S.L. See Coleman, F.C., 457 See Surman, C.A., 474
Williamson, C.E. See Smith, S.L., 136 Work, R.C. See Thomas, L.P., 474
See Zagarese, H.E., 477 Worm, B., 405, 477
Williamson, M., See Lotze, H.K., 401
Williamson, S. See Carcelles, A., 309 See Wahl, M., 475
Willis, B.L. See Babcock, R.C., 454 Worrest, R.C., 477
Willis, J.K., 476 See Häder, D.P., 462
Willis, M.J. See Simeone, A., 337 See Karanas, J.J., 465
Wilson, A. See González, J., 318 Wright, J. See Both, C., 455
Wilson, G.D.F., 172 Wyllie-Echeverria, S. See Short, F.T., 404
See Poore, G.C.B., 170 Wyrtki, K., 194, 344
Wilson, J.B. See Roberts, J.M., 472
Wilson, K.S. See Binckley, C.A., 455
Wilson, M., 172 X
Wilson, M.E.J., 194
Wilson, P.L. See See van Dalfsen, J. A., 405 Xavier, J.C. See Arata, J., 305
Wilson, S., 405
Wilson, S.G. See Gordon, H.B., 462
Wilton, K.M. See Rogers, K., 472 Y
Winant, C.D. See Jackson, G.A., 81
Wing, B.L., 88, 138 Yahia, N.D. See Bonnet, D., 455
Wing, S.R., 88, 343 Yaikin, J., 344
Winkel, W. See Both, C., 455 Yamada, S.B. See Menge, B.A., 327
Winkler, F.M. See Acuña, E., 304 Yamaguchi, H. See Tanaka, Y., 136
See Véliz, D., 342 Yamamoto, G. See Asano, E., 33
Winkler, G., 138 Yamamoto, M., 138
Winton, M. See Gregory, J.M., 462 Yamanaka, Y. See Orr, J.C., 470
Wishner, K.F., 344 Yáñez, E., 344
See Levin, L.A., 324 See Castilla, J.C., 310
Witherington, B.E., 476 See Silva, C., 337
Witman, J.D., 172 Yang, P. See Dubovik, O., 34
See Ellis, J.C., 314 See Hu, Y.-X., 34
See Fariña, J.M., 315 Yannicelli, B., 344
Wittman, K.J., 138 Yap, K.S. See Cubasch, U., 458
See Ariani, A.P., 125 Yarish, C. See Chopin, T., 311
Woebken, D. See Kuypers, M.M., 323 Yearsley, J.M. See Emmerson, M., 128
Woese, C.R., 344 Yen, A. See MacNally, R., 169
Wolanski, E., 476 Yesaki, M., 194
See Furukawa, K., 461 Yntema, C.L., 477
See Rothlisberg, P., 472 Yohe, G. See Parmesan, C., 470
533
AUTHOR INDEX
Yokoyama, M. See Takeuchi, T., 136 Zavala, P. See Castilla, J.C., 310
Yonge, C.M., 405 Zavodnik, N., 405
Yool, A. See Orr, J.C., 470 See Meinesz, A., 401
Yoon, H.S., 88 Zavoli, E. See Trombini, C., 404
Yoon, Y.H. See Ohtsuka, S., 134 Zeballos, J. See Arntz, W.E., 306
York, J.K. See Valiela, I., 405 Zeebe, R.E. See Riebesell, U., 472
Young, B. See Sayre, R., 335 Zege, E.P. See Kokhanovsky, A.A., 35
Young, C.C. See Da Silva, A.M., 312 Zein-Eldin, Z.P., 477
Ysebaert, T. See Thrush, S.F., 474 Zeiss, B. See Kroencke, I., 323
Ysla-Chee, L.A. See Narvarte, M.E., 329 Zeldis, J. See Boyd, P.W., 456
Yule, A.B. See Crisp, D.J., 312 Zenkevitch, L., 172
Yule, G.U., 172 Zentara, S.J. See Kamykowski, D., 322
Yuras, G., 344 Zerba, K. See Stephens, J.S., Jr, 86
See Morales, C.E., 328 Zertuche-Gonzalez, J.A. See Ladah, L.B., 82
See Piñones, A., 332 See Chopin, T., 311
See Ladah, L.B., 323
Zhang, J. See Keller, A.A., 465
Z Zhang, R.H. See Wang, H.J., 343
Zhang, Y., 344
Zabala, M. See Carola, M., 126 See Hutchins, D.A., 321
See Coma, R., 126 See Mantua, N.J., 468
See Delgado, O., 396 Zhao, T.X.-P., 36
See Ribes, M., 134 Zhican, T. See Rhoads, D.C., 471
Zabloudil, K. See Grove, R.S., 462 Zhivotovsky, L.A., 138
Zach, S. See Menzel, A., 469 Zimmer, M. See Orr, M., 84
Zacherl, D., 477 Zimmerman, R. See Minello, T., 402
Zacherl, D.C., 344 Zimmerman, R.C., 88, 477
Zagaglia, C.R., 477 See Kopczak, C.D., 82
Zagal, C., 344 Zipf, G.K., 172
Zagarese, H.E., 477 Zito, F. See Bonaventura, R., 455
Zagursky, G., 138 Zobell, C.E., 88
Zaitsev, Y.P., 405 Zohary, T. See Hillebrand, H., 34
Zalakevicius, M. See Lehikoinen, E., 467 Zondervan, I. See Engel, A., 460
Zamon, J.E. See Smith, S.L., 136 See Riebesell, U., 472
Zamora, S., 344 Zou, D.H. See Gao, K S., 477
Zamorano, J.H., 344 Zouhiri, S., 138
Zamzow, J.P., 477 See Dauvin, J.-C., 127
See Losey, G.S., 467 See Mouny, P., 133
Zaneveld, J.R. See Twardowski, M.S., 36 See Vallet, C., 137
Zaneveld, J.R.V., 36 Zuleta, A. See González, J., 318
See Kitchen, J.C., 35 Zuljevic, A. See Meinesz, A., 401
Zang, H. See Both, C., 455 Zuniga, O. See Ortlieb, L., 330
Zann, L.P., 477 Zust, A. See Menzel, A., 469
Zanzi, D. See Bavestrello, G., 394 Zuta, S., 344
534
SYSTEMATIC INDEX
References to articles are given in bold type, references to text pages are given in regular type.
A B
Acanthina monodon, 265, 266 Bacillariophyceae, 9
Acanthocyclus gayi, 268 Balaena mysticetus, 109
hassleri, 268 Balaenidae, 222
Acanthomysis, 104 Balaenoptera, 223
stelleri, 96, 104 acutorostrata, 223
Acartia, 215 musculus, 223
tonsa, 214, 217 physalus, 223
Acrocalanus gibber, 430 Balaenopteridae, 223
Acropora, 178, 184 Balanus laevis, 66
aspera, 185 Beggiatoa, 231, 232
Belone belone, 108
formosa, 184
Blepharipoda spinimana, 272
intermedia, 184
Bonamia ostreae, 381
muricata, 184
Brachyistius frenatus, 64
pulchra, 185, 187
Brachyramphus marmoratus, 104
Adenocystis utricularis, 276
Bryozoa, 256
Agonus cataphractus, 108
Bugula, 66
Alariaceae, 42
Alcedo pusilla pusilla, 447
Allopetrolisthes, 67 C
Alosa pseudoharengus, 108, 123
Americamysis alleni, 102, 106 Calanus, 214
bigelowi, 96, 101, 102, 106 chilensis, 214, 216, 217, 262
stucki, 102 Calidris ferruginea, 429
Anchialina agilis, 94, 95, 98, Calliclinus genicuttatus, 67
Anisotremis scapularis, 67 Calyptraea trochiformis, 67
Anomura, 263 Cancer setosus, 253, 254, 267, 268
Anous stolidus, 434 Candacia, 217
tenuirostris melanops, 445, 447 rostrata, 217
Anthozoa, 256 Carnivora, 222
Aplodactylus punctatus, 67, 241 Carpophyllum flexuosum, 445
Archaea, 303 Carukia barnesi, 432
Archaeomysis, 105 Caulerpa taxifolia, 356, 369, 370, 371, 376
Cavolinia longirostris, 440
japonica, 96, 104, 105
Centropages brachiatus, 214, 216, 217
kokuboi, 96, 104, 105
Centrostephanus rodgersii, 435
Arctocephalus australis, 222
Ceratium contortum, 213
pusillus doriferus, 412
gibberum, 213
Argopecten purpuratus, 254, 257, 263, 279, 280, 281, 292,
longipes, 29
293, 294, 296
macroceros, 213
Ascophyllum nodosum, 379
tripos, 213
Asellota, 163 Cervimunida johni, 232, 284, 285, 287
Asparagopsis armata, 66, 236, 237, 238, 241 Cetacea, 222
Asteroidea, 256 Chaetoceros, 210, 213
Atractoscion nobilis, 103 coarctatus, 213
Auchenionchus microcirrhis, 67 compressus, 213
Aulacomya ater, 236, 237, 238, 241 curvisetus, 213
Austromegabalanus, 66 dadayi, 213
psittacus, 236, 237, 238, 239 socialis, 213
Austromenidia laticlavia, 67 Charadriiformes, 222
Avicennia, 435 Charadrius bicinctus, 429
marina, 435 Cheilodactylus variegatus, 67, 241
535
SYSTEMATIC INDEX
Chelonia mydas, 427, 428, 429 Durvillaea, 274

Chionoecetes opilio, 115 antarctica, 236-240, 275
Chironex fleckerii, 431 Dynophyceae, 9
Chiton granosus, 265, 266, 267
Chlamys islandica, 263
Chlorophyceae, 9 E
Chondracanthus chamissoi, 275, 276
Ecklonia radiata, 433
Chondrus canaliculatus, 275
Eisenia, 51
Chordaceae, 42
arborea, 44, 62
Choromytilus chorus, 236, 238, 278
Embiotoca lateralis, 64, 70
Chorus giganteus, 254
Emerita, 229
Chromis chrusma, 67
analoga, 228, 229, 230, 272
Clupea harengus, 115
Emmelichthys nitidus, 431
Codium fragile, 296, 298
Engraulis encrasicholus, 108
fragile ssp. tomentosoides, 380
ringens, 218, 270, 272, 288
Coeloseris mayerii, 186
Enhydra lutris, 68
Concholepas, 278
Eretmochelys imbricata, 428
concholepas, 241, 254, 265, 266, 268, 269, 271, 272,
Erythrops elegans, 95, 98, 99, 100, 101, 106
278, 279, 282, 292, 296
erythrophthalma, 98, 101, 102, 106
Copepoda, 214
Eschrichtius robustus, 109, 122
Corallina officinalis, 236-239
Euaetideus bradyi, 217
Corallinaceae, 386
Eubalaena australis, 222, 412
Corallinales, 241
Eucalanus, 214, 216
Corycaeus typicus, 214, 217
attenuatus, 217
Cosmarestias lurida, 69
inermis, 214, 216, 217
Costariaceae, 42
subtenuis, 217
Crangon, 109
Eucampia cornuta, 210, 213
septemspinosa, 109, 110
Euchaeta, 217
Crassilabrum crassilabrum, 67
Eudyptula minor, 430
Crassostrea angulata, 383
Euphausia, 217
gigas, 292, 294, 381, 382, 383
distinguenda, 217
Crepidula, 60
eximia, 214, 217
fornicata, 381, 385, 386
mucronata, 214, 216, 217, 224, 262
Crustacea, 197
recurva, 217
Cryptophyceae, 9
tenera, 217
Cyanophyceae, 9
Euphausiacea, 214
Cymodocea nodosa, 368, 376
Euvola ziczac, 263
Cystoseira, 376, 379, 380
Exacanthomysis arctopacifica, 103
Excirolana, 229, 230
braziliensis, 228
D hirsuticauda, 228, 229
Delphinidae, 223 monodi, 228
Delphinus delphis, 223
Demospongiae, 256
Detonula pumila, 210, 213 F
Dicentrarchus labrax, 108 Fissurella, 241, 278, 292, 296
Dicrurus bracteatus carbonarius, 447 Flabellifera, 163
Dictyoneuropsis, 42 Foraminifera, 163, 256
Dictyoneurum, 42 Fucales, 379
Dinophysis rapa, 213 Fucus, 376, 377, 379, 380, 434
Diogenichthys laternatus, 271 vesiculosus 377, 378
Diomedea epomophora, 221 virsoides, 379
exulans, 221
Diplopsalis lenticula, 213
Donax, 229 G
Dosidicus gigas, 219
Doydixodon laevifrons, 67 Gadus morhua, 108, 115
Dugong dugon, 412, 428 Gastroclonium parvum, 275
536
SYSTEMATIC INDEX
Gastrosaccus psammodytes, 111 J

sanctus, 94
spinifer, 94, 95, 97, 98, 99, 106 Janus edwardsii, 436
Gelidiales, 66 Jehlius cirratus, 271
Gelidium, 241, 276
chilense, 236-239, 275
Geophyrocapsa oceanica, 431, 452 K
Girella laevifrons, 241
Kelletia kelletii, 70
Globicephala, 223
Kogia simus, 223
macrorhynchus, 223
Kogiidae, 223
melas, 223 Kyphosus analogus, 241
Glossophora kunthii, 66, 236, 237, 238, 275
Goniastrea aspera, 186, 187, 188
retiformis, 186 L
Gracilaria, 276, 293, 301
chilensis, 292, 293, 294 Labyrinthula zosterae, 372
Grampus griseus, 223 Lagenorhynchus obscurus, 223
Graus nigra, 67 Lamellibrachia, 232
Griffithsia chilensis, 66 Laminaria, 376, 380
Guinardia delicatula, 210, 213 digitata, 379
Gymnodinium, 210, 213 farlowii, 62
Gymnogongrus, 276 hyperborea, 377, 379
saccharina, 380
Laminariaceae, 42
H Laminariales, 40, 42, 241
Larosterna inca, 222
Haliotis, 264, 301 Larus belcheri, 222
discus hannai, 292, 295 dominicanus, 222, 245
laevigata, 427 modestus, 221, 222
rubra, 436 Lepidochelys olivacea, 427
rufescens, 292, 295 Leptocylindrus danicus, 213
Halophila stipulacea, 368 Leptomysis, 111
Halopteris funicularis, 236, 237, 238, 239 gracilis, 95, 98, 99, 100, 106, 111
paniculata, 66, 241 Leptonychotes weddellii, 109
Haplostylus lobatus, 94, 95, 98, 99 Lessonia, 42, 64, 236, 238, 240, 241, 242, 244, 246, 274,
lobatus var. armata, 94 276
normani, 94, 95, 98, 99 nigrescens, 236-242, 254, 257-259, 261, 275, 277, 296
Helcogrammoides cunninghami, 67 trabeculata, 66, 236-242, 245, 257, 260, 275, 277, 296
Heliaster, 71, 72 Lessoniaceae, 42
helianthus, 241, 259, 261 Lessoniopsis, 42
Hemilutjanus macrophthalmos, 67 Leucocarbo bougainvilli, 222, 225
Heterocarpus reedi, 284 -287 Libidoclaea granaria, 272
Heteroclinus, 434 Limnoria chilensis, 254
Hildenbrandia, 66 Liparis liparis, 108
Hippoglossina macrops, 272 Lissodelphis peronii, 223
Homalaspis plana, 268 Lithophaga lithophaga, 379, 380
Hypoleucos brasiliensis, 222 Lithothamnion corrallioides, 383, 386
Hypsoblennius sordidus, 67 glaciale, 383
Lobodon carcinophagus, 109
Loligo forbesi, 432
I Lontra felina, 69, 222, 224, 241, 296
Loxechinus albus, 65, 69, 241, 260, 279, 283, 292, 296
Iiella ohshimai, 96, 104, 105 Lucinidae, 232
Isacia conceptionis, 67, 278 Luidia, 69, 71, 72
Isopoda, 256 magellanica, 241, 261
537
SYSTEMATIC INDEX
M Neotrypaea uncinata, 272

Nephthys incisa, 145
Macrocystis, 39-88, 236, 238, 242, 274, 283, 428 impressa, 228, 229
angustifolia, 42, 43, 44, 47, 51, 63 Nereocystis, 42, 72
integrifolia, 42-46, 47, 51, 53, 54, 57, 60, 63, 64, 66, Nipponomysis ornata, 96, 105
67, 236, 237, 239, 240, 242, 243, 244, 259, 260, toriumi, 108
261, 275, 296 Noctiluca, 12
laevis, 42, 43, 44, 46, 63 scintillans, 431
pyrifera, 42-47, 53, 54, 56, 60, 63, 64, 236, 237, 240, Nothochthamalus scabrosus, 265, 266
257, 261, 275 Notobalanus flosculus, 251
Macruronus novaezelandiae, 434 Nucula proxima, 145
Maldane sarsi, 164
Maurolicus parvipinnis, 272
Mazzaella, 276 O
laminarioides, 254, 275
Megaptera novaeangliae, 412, 223 Oceanodroma tethys, 222
Merlangius merlangus, 108 tethys kelsalli, 296
Merluccius gayi, 219, 272, 284 Ocypode gaudichaudii, 228
gayi gayi, 253 Oithona similis, 214, 217
merluccius, 108 Oncaea conifera, 214, 217
Mesodesma donacium, 228, 229, 230 230, 254, 279, 280, Ophiaycthis kroyerii, 67
281, 292, 294 Ophiura sarsi, 164
Mesophyllum, 66 Orchestoidea tuberculata, 228-230
Mesopodopsis slabberi, 95, 97, 98, 101, 103, 106, 111 Orcinus orca, 223, 224
wooldridgei, 97, 111 Ornithocercus magnificus, 213
Metamysidopsis elongata, 96, 103, 115 Ostrea edulis, 380, 381, 382, 383, 390
Meyenaster, 69, 71, 72 Otaria flavescens, 219, 222, 224, 241, 278
gelatinosus, 241 juvata, 278
Mitrella unifasciata, 67 Otariidae, 222
Mollusca, 256 Ovalipes trimaculatus, 268
Montastraea faveolata, 188
Montemaria horridula, 238
Montipora, 184 P
digitata, 184
ramosa, 185 Pachyptila desolata, 221
Morus serrator, 428 Pagurus, 67, 272
Mugil cephalus, 67 Palythoa, 187
Mustelidae, 222 Pandalus borealis, 115
Mya arenaria, 356 Panulirus, 68
Mycedium elephantotus, 187 cygnus, 412, 435
Mysidacea, 90 Paracalanus, 214
Mysideis parva, 96, 99, 100, 101 parvus, 214, 217
Mysidopsis bigelowi, 101, 102, 106, 112 Paralabrax humeralis, 67
Mysis relicta, 110 Paralichthys californicus, 108
stenolepis, 111 olivaceus, 105
Mytilus edulis, 386 Paraprionospio pinnata, 232, 234, 262
trossulus, 264 Pecten, 56, 63
maximus, 385
Pelagophycus, 42
N porra, 42
Pelecaniformes, 222
Nassarius gayi, 67 Pelecanoides garnotii, 222, 296
Natator depressus, 412 Pelecanus thagus, 222
Nematoscelis flexipes, 217 Penaeus merguiensis, 412, 446
megalops, 217 Perkinsus, 427
Neomysis americana, 111, 113, 117, 123, 124 Perumytilus purpuratus, 236-239, 251, 265, 266
kadiakensis, 96, 103, 106, 108, 112, 117 Petrolisthes, 263, 264
rayii, 96, 103, 104 granulosus, 263, 264
Neophoca cinerea, 412 laevigatus, 263, 264
538
SYSTEMATIC INDEX
tuberculatus, 263, 264 R

tuberculosus, 263, 264
violaceus, 263, 264 Raja clavata, 100
Phaeophyceae, 40, 275 Rexea solandri, 434
Phaeophyta, 256 Rhincalanus nasutus, 272
Phalerisida maculata, 228, 229 Rhizophora apiculata, 441, 444
Phocoena spinipinnis, 223 mangle, 438, 446
Phocoenidae, 223 Rhizosolenia, 210
Phragmatopoma, 66 imbricata, 213
moerchi, 236-239, 241 Rhodophyta, 256, 274
Phyllophora, 379 Rhodymenia skottsbergii, 238
Phyllospora comosa, 428 Rhynchocinetes typus, 283
Phymatolithon calcareum, 383, 385, 386 Ruppia cirrhosa, 368
maritima, 368
Physeter macrocephalus, 223
Physeteridae, 223
Pinguipes chilensis, 67, 241
Pinnaxodes chilensis, 267
S
Pisoides edwardsi, 268 Sabellaria, 391
Platichthys flesus, 108, 432 alveolata, 386, 387
Pleuromamma gracilis, 217 spinulosa, 386, 387
Pleuroncodes monodon, 232, 272, 273, 284, 285 Sardinops sagax, 218, 288, 429, 449
Pleuronectes platessa, 108 Sargassum, 376, 379
Pocillopora, 184 hornschuchii, 379
damicornis, 184 muticum, 379
Polychaeta, 256 Scartichthys viridis, 67
Pomatoschistus lozanoi, 108 Schistomysis, 98
minutus, 108 kervillei, 95, 98, 106
Porcellanidae, 263 ornata, 95, 98-101, 106
Porifera, 66, 256 spiritus, 95, 97, 98, 101, 106
Porites, 184, 185, 187, 189, 439 Schroederichthys chilensis, 241
lutea, 184, 186 Scolecithrichella bradyi, 217
(synaraea) rusi, 184 Scomber japonicus, 288
Porphyra, 276 Scurria, 67
Posidonia australis, 429 scurra, 258
oceanica, 368, 369, 370, 371, 373, 375, 376 Scytothamnus fasciculatus, 276
Postelsia, 42 Semicossyphus, 68
Potamocorbula amurensis, 109 maculatus, 241
Prisogaster niger, 67 Semimytilus algosus, 236, 237, 238, 251
Sepia officinalis, 109
Procellaria aequinoctialis, 221
Siriella japonica, 108
Procellariiformes, 222
Skeletonema costatum, 213, 446
Prochlorococcus, 430
Solemyidae, 232
Prolatilus jugularis, 67
Sphenisciformes, 222
Prorocentrum micans, 213
Spheniscus humboldti, 221, 222, 296
Psammobatis scobina, 241
magellanicus, 222
Psammocora digitifera, 187 Spionidae, 66
Pseudochordaceae, 42 Sterna bergii, 413
Pseudo-nitzschia cf. delicatissima, 213 fuscata, 434, 445
pseudoseriata, 213 lorata, 222
Pseudorca crassidens, 223 Stichaster, 71,72
Pterodroma externa, 221 striatus, 241, 259
Pterygophora, 49, 50, 51, 53 Stictocarbo gaimardi, 222
californica, 62 Strangomera bentincki, 272, 288
Puffinus pacificus, 428 Strongylocentrotus, 68
tenuirostris, 413, 428, franciscanus, 65
Pycnopodia, 68 purpuratus, 65
Pyura chilensis, 66, 236-238, 241, 259 Stylocheiron affinis, 217
praeputialis, 236-240 Subeucalanus crassus, 217
539
SYSTEMATIC INDEX
Sula variegata, 222, 225 Tursiops truncatus, 223, 296

Symbiodinium, 187, 419 Tylos spinulosus, 228, 229, 230
Synechococcus, 430
U
T Undaria pinnatifida, 379
Tagelus dombeii, 292, 294 Uria aalge, 104
Taliepus, 67
Tegula, 241, 264
atra, 67, 278 V
tridentata, 67 Vermetidae, 66
Teleostei, 256 Vesicomyidae, 232
Tetrapygus, 69, 72
niger, 67, 69, 71, 241, 260, 261
Thalassarche bulleri, 221 X
eremita, 221
Thalassiosira, 210, 213
Thioploca, 231 Xantochorus cassidiformis, 67
Thunnus maccoyii, 431 Xenacanthomysis pseudomacropsis, 103, 104
Thyasiridae, 232
Trachurus declivis, 431, 434
murphyi, 67, 219, 220, 253, 278, 288
Z
symmetricus, 288 Ziphiidae, 223
Tricolia mcleani, 67 Ziphius cavirostris, 223
Trigla lucerna, 108 Zostera, 99, 373
Triphoturus oculeus, 271 marina, 368-376, 446
Trisopterus luscus, 108 noltii, 368, 370, 371
540
SUBJECT INDEX
References to complete articles are given in bold type, references to sections of articles are given
in italics, references to pages are given in regular type.
A Cavitation, 55
CHALLENGER, 143, 146, 164, 197
Absorption coefficient, 4, 10, 31, 37 Charles Darwin, 40, 173, 201
Absorption, 1, 4, 5, 6, 7, 10, 12, 13, 14, 16–19, 20, 23, Chile Coastal Current, 202, 203, 204
25–28, 30, 31, 32, 37 Chlorophyll a, 1, 210, 250, 265, 267, 269, 271, 273, 291
Acoustic Doppler current profiler (ADCP), 90 Chlorophyll, 207, 212, 213, 224, 270, 437
Adriatic Sea, 351, 354, 355, 365, 370, 375, 380, 386, 387 Climate change and Australian marine life, 407–478
Advection, 59 Australian marine biodiversity, 411–413
Aeolian dust, 411 climate change projections for Australia, 413–419
Aerosols, 413, 416, 441 CO2, pH and calcium carbonate saturation state,
Altimeter, 181, 182 415–416
Analytical species accumulation (ASA), 160 mixed layer depth and stratification, 415
Andaman Sea, coral reefs, 173–194 ocean currents, 415
Apogamy, 49 ocean temperature, 414
Aquaculture, 242, 273, 279, 280, 284, 291, 292–295, 296, precipitation and storms, 417–418
298, 301, 302, 354, 356, 363, 367, 371, 381, sea level, 418–419
385, 388, 390, 392, 409, 449 solar radiation, 416
Areas of Special Scientific Interest, (ASSI), 368, 376, 389 winds, 415
Attenuation, 1, 5, 6, 10, 12, 14, 16–19, 20, 23, 25–32 climate impacts on Australian marine life, 419–447
CO2, pH and aragonate saturation state, 438–441
currents, 435–436
B mixed layer depth and stratification, 436–438
precipitation and storms, 443–446
Backscatter(ing), 5, 6, 10, 14, 16–26, 30, 31, 32, 37, 38, sea level, 446–447
114, 117, 120, 121 solar radiation, 441–443
Bacteria, 2, 7, 8, 31, 50, 187, 196, 211, 212, 231, 232, 234, temperature, 419–433
245, 270, 440, 442 winds, 433–434
Baltic Sea, 351, 353, 354, 357, 359, 360, 367, 368, 370, community impacts, 447–448
371, 374, 377, 380, 388 critical knowledge gaps and a way forward, 452–453
Benthic boundary layer, 52, 55, 57, 59, 62 non-climate stressors, 448–449
Benthic-pelagic coupling, 90, 112, 198, 299, 300 summary, 449–452
Biodiversity informatics, 140, 165 temperate Australia, 451–452
Biodiversity, 62, 140, 146, 148, 164, 197, 201, 214–215, tropical and subtropical Australia, 451
236, 240, 295–299, 301, 302, 407, 411–413, Climate change, 62, 71–73, 277, 278, 299, 303
431, 436, 445, 453 Climate models, 407, 413, 414, 415, 416, 417, 418, 419,
Biogeography, 140, 146, 164, 174–179, 196, 200, 214–215, 426, 428, 434, 478
252, 254, 255–262, 264, 273, 274, 276, 277, Coastal defence, 355, 367, 379, 388, 390
304 development, 345, 349, 383, 388, 390, 391
Birds Directive, 356, 358, 368, 376, 391 erosion, 354, 367, 387
Black Sea, 98, 351, 358, 359, 360, 368, 378, 379, 380, 382, flooding, 354 355
383 Coastal marine habitats in Europe, loss, status, trends,
Bleaching of corals, 183, 188, 189, 407, 419, 421, 423, 345–405
426, 427, 428, 441, 450, 451, 452 conclusions, 393
Bonn Convention, 356, 357, 388 definitions, 347–348
Bottom-up processes, 70, 198, 218, 219, 259, 261, 265 general European context, 348–360
Brooding, 247, 249, 262, 267–268 EU coastal policies and trans-national agreements,
356–360
threats to coastlines, 349–356
C habitats, 360–389
biogenic formations, 386–387
C:N ratios, 112 macroalgal beds, 376–380
Calcification, 416, 432, 438, 439, 441 maerls, 383–386
Carbon budget, 54 oyster reefs, 380–383
541
SUBJECT INDEX
seagrass meadows, 368–376 traditional approaches, 147–152

sedimentary habitats, 387–389 abundance distributions, 147–148
wetlands, salt marshes, 360–368 information theory and Shannon’s H′, 151–152
recommendations for conservation and management, Simpson’s λ, 149–151
391–393 species richness and its rarefaction, 148–149
Colonisation, 58, 59, 60 Diversity, 200, 215, 221, 230, 231, 232, 234, 237, 241, 252,
Community structure, 92, 107, 111, 425, 442, 443, 445, 448 256, 258, 259, 274, 295, 296, 301, 302, 411,
Competition, 39, 40, 44, 46, 50, 54, 56, 57, 68 412, 413, 425, 437, 445, 447
Conservation and management, 391–393 DNA, 43, 73
Conservation, 139, 140, 141, 145, 152, 154, 157, 164, 165, Downwelling, 202, 249, 270
222, 223, 227, 242–244, 250, 265, 295–299,
Dredging, 173, 189
300–301, 302, 303, 304
Drift, kelp, 62, 64, 65, 68, 71, 228, 244,
Coral reefs of the Andaman Sea, 173–194
biological characteristics of coral reefs, 184–188
physiological attributes of reef corals, 186–188
reef type and coral diversity, 184–186
E
conclusions, 190 Earthquakes, 173, 176, 189
geological history, sea floor topography and related
East Australian Current (EAC), 407, 409, 414, 415, 419,
coral biogeography, 174–179
422, 428, 431, 435, 436, 449, 452
natural and human influences, 188–190
Echo sounders, 90
physical influences affecting coral reefs, 179–184
Ecology of giant kelp, 39–88
oceanography, 179–181
ecology of communities, 62–73
sea temperatures and salinity, 182–183
sedimentation, 183–184 community consequences of climate change and
tidal influences and sea-level fluctuations, 181–182 kelp forest exploitation, 71–73
Coral reefs, 409, 411, 412, 416, 419, 421, 423, 424, 428, Macrocystis as a foundation species, 62–65
435, 439, 441, 442, 444, 445, 449, 450, 451 trophic interactions and food webs, 65–71
Coral Sea, 409, 410 organismal biology, 42–54
CORINE project, 351, 354, 361 distribution, 44–47
Coulter counter, 10 evolutionary history, 42–44
growth, productivity and reproduction, 50–54
life history, 47–50
D population biology, 54–62
demography and population cycles, 61–62
Deep sea, 411, 437, 438, 441 dispersal, recruitment, population connectivity,
Deep-sea diversity, 163–164 57–61
Deforestation, kelp, 55, 64, 65, 72 stage- and size-specific mortality, 54–57
Detritus, 7, 8, 89, 107, 110, 111, 115, 245, 246, 247
Ecosystem engineers, 236–238, 238–240
Dispersal, 39, 40, 54, 57–61, 196, 198, 199, 236, 241, 244,
Ecosystem-based management, 392
245, 247–251, 252, 254, 255, 258, 269, 296,
Ekman transport, 201, 204, 260, 273
298, 300, 303, 304, 412, 435
El Niño (EN), 56, 70, 71, 72, 196, 198, 199, 200, 204, 211,
Disruptive fishing techniques, 353, 367, 379
212, 215, 216, 217, 219, 225, 230, 235, 236,
Diversity estimations in sedimentary benthic communities,
242, 243, 244, 250, 254, 257, 258–262, 263,
139–172
273, 274, 276, 278, 280, 281, 282, 291, 293,
conclusions, 164–166
294, 304, 420, 431, 435
examining large-scale patterns, 161–164
deep-sea diversity, 163–164 El Nino-Southern Oscillation (ENSO), 51, 52, 57, 59, 61,
global latitude gradient, 161–162 62, 71, 72, 196, 198, 204, 206, 216, 218,
history, 142–146 224–225, 230, 234, 242, 245, 253, 257, 258,
end of Petersen era, 144–145 259, 260, 262, 265, 274, 288, 292, 293, 294,
Forbes zones to Petersen communities, 142–144 299, 431, 435
new era, 145–146 Endemic species, 214, 221, 223, 224, 225, 232, 273
influx of indices, 146–152 Endemism, 411, 412, 427
newer developments, 152–161 Energy budgets, 91
extrapolation, 157–158 Eutrophication, 351, 353, 367, 371, 373, 375, 379, 385, 386
progress in measurement, 158–161 Evolutionary history, Macrocystis, 42–44
surrogates, 152–154 Exploitation, 39, 62, 69, 71–73
taxonomic diversity, 154–157 Extrapolation, in diversity, 139, 157–158, 163, 164, 165
542
SUBJECT INDEX
F regionally abundant mysids and their migration habits,

97–108
Fisheries, 196, 197, 201, 206, 207, 219, 220, 221, 224, 226, European shelves, 97–101
245, 278–284, 284–288, 288–292, 295, 296, mysid ‘umwelt’, 105–108
299, 353, 357, 358, 371, 381, 383, 386, 388 northeast Pacific shelves, 103–104
Fisher’s α, 147–148 northwest Atlantic shelves, 101–103
Fishing, 412, 448, 452, 453 northwest Pacific shelves, 104–105
Fjords, 100, 101, 105, 106, 113, 114, 123 other regions, 105–108
Fluorine cycle, 92 Habitat degradation, 346, 347, 348, 360, 361 363, 388, 391
Fluxes, horizontal, 89, 106, 107, 113, 117 fragmentation, 346, 347, 348, 368, 393
Food webs, 40, 62, 65–71, 89, 91, 107, 108–112, 113, 115, loss, Europe, 345–405
196, 197, 210–214, 218, 219, 220, 240, 261, modification, 346, 349, 368, 389
269, 270 Habitats Directive, 347, 356, 358, 360, 367, 368, 376, 380,
Forbes zones, 142–144 383, 386, 389, 390, 391
Founder effects, 44 Herbivores, 241, 242, 245, 265, 266, 267
Humboldt Current System, 195–344
aquaculture, 292–295
G artisanal benthic fisheries, 278–284
case studies, 280–283
Gametogenesis, 48, 49
Chilean abalone fishery, 282–283
Gene flow, 40, 42, 43, 44, 53, 73, 247, 252, 253, 254, 258
scallop fishery, 280
Genetic diversity, 252, 253, 254, 258
sea urchin fishery, 283
relatedness, 43
surf clam fishery, 280–282
Geometric optics region (GO), 8, 12, 13, 14, 16
resource dynamics and management, 283–284
Georges Bank, 101, 102, 109
bio-economic aspects of industrial crustacean fisheries,
Giant kelp, ecology, 39–88
284–288
GIS, 298
basic biology of nylon shrimp and squat lobsters,
Glaciation, 176, 178
285–286
Global Coral Reef Monitoring Network (GCRMN), 174
biology and stock estimates, 286
Global warming, 173, 187, 189, 199, 208, 264, 367, 415,
crustacean fisheries and economic considerations,
418, 435, 450
286–288
Grazing, 40, 46, 55, 56, 63, 64, 65, 66, 68, 69, 71
biogeography, 255–262
Great Barrier Reef, 407, 408, 410, 411, 412, 419, 421, 422,
El Niño-La Niña in coastal marine communities,
424–427, 429, 430, 434, 439, 440, 444, 445,
258–262
447, 451
large-scale patterns, 255–257
Greenhouse gases, 407, 409, 413, 416, 449
role of past and present processes, 257–258
brief history of exploitation of natural resources,
277–278
H first coastal settlers, 277–278
Habitat alteration, 112–113 coastal oceanography, 205–208
conversion, 346, 347, 363, 365, 367, 390 conclusions, 303–304
Habitat coupling by marine mysids, 89–138 conservation of marine biodiversity and Marine
an appreciation of mysids, 108–122 Protected Areas, 295–299
mysid importance in the coastal marine economy, dominant primary producers and their role in the pelagic
108–115 food web, 211–214
dominating the holoplankton, 113–115 processes affecting primary production and export
food-web roles, 108–112 108–112 processes, 213–214
habitat alteration and coupling, 112–113 export and import processes, 244–246
lending trophic and dynamic stability, 115 between benthic habitats, 245–246
reasons why mysids have been under-appreciated, between environments, 245
115–122 between realms, 244–245
modelling difficulty, 117–122 frequency and intensity, 246
sampling problems, 115–117 fish consumers, 218–219
challenges and opportunities, 122–124 history of research, 200–211
methods of data collection, 94–96 first reports on a cool current in the SE Pacific, 200
migratory capabilities, schooling and consequences, from natural history to ecology to marine
92–94 conservation, 200–201
543
SUBJECT INDEX
intertidal and subtidal hard-bottom communities, propagule supply, dispersal and recruitment variability,
235–240 246–252
habitat-forming species on hard bottoms, 236 dispersal, 248–249
macrofauna associated with ecosystem engineers on methodological approaches to the study of
hard bottoms, 238–240 dispersal, 247–248
spatial and temporal dynamics of ecosystem patterns of recruitment and benthic communities,
engineers, 236–238 250–252
kelp forests, 240–244 settlement, 249–250
conservation and human activities, 242–244 reproductive patterns of selected marine invertebrates,
population dynamics and spatial distribution, 265–268
241–242 primary productivity and gonad production,
the community, 241 265–267
larval life, 268–273 temperature and brooding requirements, 267–268
behavioural traits, 268–269 sandy beaches, 227–230
feeding and larval food environment, 269–271 seabirds and marine mammals, 219–227
upwelling and larval transport processes, 271–273 effects of ENSO on seabirds, 224–225
life history adaptations of macroalgae, 273–277 food resources or breeding sites, 225–227
environmental gradients and strategies of marine mammals, 224
resistance, recovery and recolonisation, 274 seabirds, 219–224
enhanced solar radiation, 277 subtidal soft-bottom communities, 230–235
physiological and morpho-functional
latitudinal and bathymetric patterns, 232–234
adaptations, 276–277
temporal patterns of variability in shelf
temperature tolerance, 276
communities, 234–235
oceanographic conditions in the SE Pacific, 201–205
zonation patterns, 230–232
outlook, long-term research vision and future research
water column chemistry, 208–214
frontiers, 299–303
macro- and micronutrient distribution, 210–211
frontier research opportunities, 303
oxygen distribution and relevance in the organic
long-term scientific vision, 302–303
carbon remineralization, 209–210
effects of future climate change, 303
zooplankton consumers, 214–217
high-sea conservation policy, 302
precautionary and integrative ecosystem biogeographic and biodiversity issues, 214–215
management, 302 future perspectives in zooplankton research,
slope, deep-sea and abyssal ecosystems, 302 217–218
technological research on deep-sea hydrates, coastal upwelling in northern Chile, 215
302–303 oxygen minimum zone, 215–216
short/mid-term scientific outlook, 299–302 life cycles and population dynamics, 216–217
coastal network system for marine conservation Hydrothermal vents, 153
management, 301 Hyperbenthos, 92
inshore and offshore oxygen minimum
ecosystems, 299–300
integrative and adaptive resource management I
approach to fisheries, 300
marine molecular biology, 301 Index of refraction, 3, 6–7, 11, 12, 13, 27, 29, 32, 37
marine non-indigenous species, 301 Indicator species, 140, 145, 152, 153
nearshore coastal oceanography and benthic- Individual-based models (IBM), 92, 123, 124
pelagic coupling, 300 Information theory, 146, 151–152
novel approaches in coastal mariculture, 301 Inherent optical properties (IOPs) of non-spherical marine
ocean-atmosphere interactions and offshore particles, 1–38
oceanography, 299 bulk IOPs, 3–10
training of Chilean marine taxonomists, 301–302 characteristics of particles affecting their optical
pelagic fisheries and fisheries management, 288–292 properties, 6–10
history of the catches, 288–289 index of refraction, 6–7
management of pelagic fisheries, 291–292 orientation, 10
relationships with oceanographic variations, shape, 8–10
289–291 size, 7–8
physiological adaptations of marine invertebrates, definitions, 3–6
262–265 effects of particle shape on IOPs, 29–30
population connectivity, 252–255 IOPs of monodispersions of randomly oriented
population connectivity studies, 253–255 spheroids, 13–19
544
SUBJECT INDEX
attenuation, absorption, and scattering: efficiency M

factors and biases, 16–19
exact and approximate methods, 13–14 Macroalgae, 412, 420, 422, 423, 424, 427, 428, 436, 439,
results: IOPs of a monodispersion, 14–16 441, 445, 446
Macroalgal beds, 376–380, 389, 390
volume scattering function, 14–16
Maerls, 383–386, 390, 391
optical properties of polydispersions, 19–29
Mangroves, 346, 376, 409, 411, 420, 422, 423, 424, 425,
obtaining the IOPs of polydispersions of particles, 428, 429, 435, 438, 441, 443–447, 449, 451
19–25 Marine Protected Areas (MPA), 197, 244, 295–299, 359,
results for polydispersions, 25–29 360, 376, 389, 391, 392
optical regimes, 11–13 Mediterranean Sea, 94, 98–102, 113, 350, 351, 353, 354, 356,
geometric optics region, 12 358, 359, 360, 364, 367, 368, 369, 371, 372
Rayleigh region, 11–12 Meiofauna, 110
Meroplankton, 112, 113, 432
Rayleigh-Gans-Debye and the van de Hulst regions,
Meta-analysis, 161, 162, 165
12–13
Metapopulations, 60, 61
summary and future prospect, 30–33
Microbial loop, 210, 437
Integrated Coastal Zone Management (ICZM), 392 Microphotometry, 50
Intergovernmental Panel on Climate Change (IPCC), 183, Mie theory, 2, 13, 29, 32
355, 407, 411, 414, 418 Migration, 89, 90, 91, 92–94, 97–108, 109, 110, 112, 113,
Internal waves, 181 117–120, 122, 123, 124, 216, 224, 253, 254,
International Union for Conservation of Nature and 259, 261, 285, 287, 291, 296, 408, 421, 424,
Resources (IUCN), 174 429, 432, 435, 436, 446
Intertidal zone, 44, 46, 47, 53, 72, 97, 173, 184, 185, 188, diel vertical (DVM), 216, 219, 220, 262, 269, 272
228, 229, 230, 235, 238, 240, 244, 246, 251, vertical, 90, 92, 93, 97, 100, 102, 103, 105, 109, 111,
263, 264, 265, 280, 351, 360, 363, 368, 376, 112, 123, 124, 214, 215, 272, 303
384, 387, 388, 392, 411, 419, 427, 443, 444, Missing backscattering enigma, 2
446 Modelling, 89, 117–122, 123, 205, 218, 225, 300
Molecular biology, 299, 301
Introduced species, 356, 361, 390
Monoclonal antibodies, 50
Invasive species, 367, 379, 389
Monodispersions, 1, 10, 13–19, 21, 25, 29, 30, 32
Island biogeography theory, 63 Monsoon, 179, 180, 181, 183, 184, 410, 443
Mortality, stage and size specific, 54–57
Mudflats, 347, 360, 387–389, 390
K
Kattegat, 351, 374, 380 N
Kelp, 41, 420, 433, 442, 445, 452
forests, 39, 40, 58, 61–65, 68, 70, 71–73, 238, 240–244, NATURA 2000, 356, 358, 359, 360, 373, 375, 376, 379,
245, 246, 428, 449 380, 381, 382, 384, 386
Nitrogen budget, 54
giant, ecology of, 39–88
North Sea, 98, 99, 351, 353, 357, 360, 374, 381, 382, 384,
Keystone species, 447, 448
388, 420, 421, 431, 432, 433
Nutrients, 196, 208, 209–210, 230, 240, 241, 242, 244, 245,
246, 247, 259, 260, 274
L
La Niña (LN), 56, 61, 62, 70, 71, 204, 214, 216, 225, 234,
258–262, 280
O
Land reclamation, 173, 189, 345, 350, 351, 365, 366, 375, Ocean Circulation and Climate Advanced Model
385 (OCCAM), 179
Leeuwin Current, 409, 410, 412, 415, 422, 431, 435, 452 Oil spills, 73
Life history, Macrocystis, 39, 40, 42, 47–50, 57, 59, 61, 62 Optical sensors, 1, 2
Overfishing, 345, 349, 381, 390, 434
Limitation, light, 60
Oxygen minimum zone (OMZ), 196, 208, 214, 215–216,
nutrient, 52, 55, 57
217–220, 230, 231, 232, 235, 247, 258, 285,
Logarithmic series, 146, 150 299, 301, 303
Log-normal distribution, 7, 8, 147–148, 150, 164 Oyster reefs, 345, 346, 347, 349, 378, 380–383, 389, 390
Log-series distribution, 147–148, 150, 162, 164 Ozone layer, 416
545
SUBJECT INDEX
P Sandy beaches, 227-230, 235, 246, 247, 256, 294

Sandflats, 376, 387-389, 390
Pacific Decadal Oscillation (PDO), 71 Satellites, 181, 184, 204, 207, 250, 271, 290, 416
Paramonov method, 13, 14, 17, 18, 30 Scattering, 1, 2, 4-7, 9, 10, 12-18, 20-23, 25-32, 37, 38
Particulate size distribution (PSD), 7, 8, 21, 32, 37 Schooling, 89, 91, 92-94, 97, 103, 107, 111, 112, 115, 116,
Peru Coastal Current (PCC), 202, 203 117, 122, 124
Petersen communities, 142, 145, 146, 152 Sea level, 173, 176, 177, 179, 181-182, 188, 189, 355, 356,
Phenology, 274, 275, 411, 419, 429–430, 432, 433, 434, 363, 367, 387, 388, 390, 407, 411, 413, 418-
437, 445–446, 449, 453 419, 425, 432, 446-447, 449, 450, 451
Pheromones, 48, 49 Sea-level anomalies, 173, 181
Photosynthesis, 42, 49, 51, 54, 59, 73, 267, 277, 438–442, 446 Sea surface temperature (SST), 204, 206, 207, 224, 225,
Photosynthetically active radiation (PAR), 44, 48, 49 226, 255, 261, 289, 290, 291, 294
Phytoplankton, 2, 3, 7, 8, 9, 12, 14, 17, 18, 20, 23, 24, 29, Seagrasses, 409, 411, 412, 419, 420, 422-425, 427, 428,
31, 32, 106, 109–113, 210, 212, 213, 214, 219, 429, 434, 438, 441, 442, 445, 446, 449, 451
220, 245, 269, 270, 271, 375, 410, 411, 420, Seagrass meadows, 345, 347, 349, 368-376, 390
422, 425, 430, 432, 436–440, 442–447 Sediment dynamics, 89, 91
Pneumatocysts, 47, 64 Sedimentary habitats, 387-389
Polarised light, 93, 112, 115, 122 Sedimentation, 7, 173, 183-184, 410, 444, 447
Pollution, 242, 269, 274, 295, 300, 301, 345, 346, 349, 351, Seston, 89
353, 356, 357, 358, 367, 375, 377, 383, 388, Settlement, 198, 234, 241, 249-250, 261, 268, 269, 272,
390, 392, 448, 449, 451 277, 287
Polydispersions,1, 10, 13, 14, 19–29, 30, 32 Sex determination, 47
Population biology, Macrocystis, 54–62 Shannon's index, H′, 139, 145, 151-152, 153, 162, 163, 165
connectivity, 57–61, 196, 248, 252–255, 261, 262, 298, Shifting baselines, 345, 391
304 Shoaling, 104, 107, 118
growth, 350, 351 Simpson's index, λ, 139, 146, 149-151, 154, 155, 156, 162,
Primary production (PP), 196, 197, 198, 205, 207, 208, 163, 165
209, 210, 211, 212-214, 234, 259, 265, 267, Solar radiation, 407, 411, 413, 416, 418, 441-443
270, 271, 274, 282, 303 Solitons, 181
Protection initiatives, 357, 358 Special Areas of Conservation (SAC), 356, 358, 368, 376,
Pycnocline, 181 386
Special Protection Area (SPA), 351, 356, 358
Species richness, 139, 141, 143, 144, 145, 148-149, 151,
R 152, 153, 155-163, 165
Stable isotopes, 70
Ramsar Convention on the Conservation of Wetlands, 356, Statoliths, 92, 93
357, 367, 388 Storm waves, 59, 65, 71
Rarefaction, 139, 145, 146, 148-149, 163, 165 Storms, 407, 411, 413, 417-418, 424, 425, 429, 443-446,
Rayleigh region (RAY), 8, 11-12, 19 448, 451
Rayleigh-Gans-Debye region (RGD), 8, 12-13 Surf zone, 52, 89, 97, 102-107, 111
Recombinant DNA, 43 Surrogates, for biodiversity, 152-154, 156, 160, 162
Recruitment, 39, 40, 47-50, 53, 54, 57-61, 62, 63, 68, 73,
197, 199, 217, 218, 229, 230, 234, 236, 238,
239, 241, 247-250, 250-251, 254, 258, 259, T
261, 269, 271, 274, 280-284, 286, 293, 304,
422, 434, 435 Tasman Sea, 407, 408, 409, 414, 431, 452
windows, 48 Taxonomic Distinctness, 154, 156, 157
Remote sensing, 44 Diversity, 154, 156
Ribosomal RNA genes, 187 Tectonics, 174, 175, 176, 177, 179
Rio Convention on Biological Diversity (CBD), 356, 357, Thermocline, 181, 182, 487
367, 376, 391 Thorson's rule, 144
Rocky shores, 420, 422, 423, 427 Tidal transport, 272
T-matrix method, 13, 14, 16, 17, 18, 21, 30, 31
Top-down consumption, 70, 71
S processes, 198, 212, 219, 250, 261
Total Allowable Catch (TAC), 284, 286, 291
Salt marshes, 346-349, 352, 354, 357, 360-368, 390 Tourism, 206, 221, 284, 295, 296, 298, 299, 302, 349, 354,
Sampling efficiency, 97, 104, 116 358, 367, 371, 451, 453
546
SUBJECT INDEX
Trawling, 345, 368, 376, 379, 383, 386, 390 Volcano, 175, 176
Trophic focusing, 107 Volume scattering function (VSF), 4, 5, 7, 12, 14-16, 37
interactions, 65-71
Tsunamis, 188, 189
W
U Wadden Sea, 348, 350, 351, 357, 359, 360, 365, 368, 373,
378, 379, 381, 382, 383, 386, 387, 391
Ultraviolet radiation (UV), 44, 199, 269, 277, 413, 416, Wasting disease, 372, 373, 374
423, 441, 442, 443 Waves, 44, 46, 51, 52, 54, 55, 57, 68
Umwelt, of mysids, 105-108, 117 West Wind Drift (WWD), 201, 202, 203
United Nations Environment Programme (UNEP), 174, 180 Wetlands, 345, 346, 356, 357, 359, 360-368, 389, 390, 391
Upwelling, 181, 182, 187, 196-199, 201, 202, 204-214, Winds, 407, 411, 413, 414, 415, 417, 422, 433-434, 436,
217, 218, 221, 224, 225, 230, 231, 234, 236, 437
242, 244, 245, 247, 248, 249, 250, 252, 257, World Wildlife Fund (WWF), 351
258, 260, 261, 262, 265-274, 280, 282, 289,
290, 299, 301, 303, 304, 410, 415, 433
Z
V Zooplankton, 7, 8, 12, 14, 90, 101, 106, 109-113, 117, 123,
196, 197, 198, 211, 212, 214-218, 219, 220,
Van de Hulst region (VDH), 8, 12-13 410, 420-423, 430, 432, 433, 434, 437, 440,
Vinogradov's ladder, 90, 123 443, 447, 452, 453
Viruses, 7, 8, 30 Zooxanthellae, 186, 188
547

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fm Page 16 Thursday, May 10, 2007 8:38 PM

WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS

Attenuation, absorption and scattering: efficiency factors and biases

INHERENT OPTICAL PROPERTIES OF NON-SPHERICAL MARINE-LIKE PARTICLES

Phase shift parameter ρ Phase shift parameter ρ

0.5 1 2 5 10 20 50 100 200 Qc 0.5 1 2 5 10 20 50 100 200 Qa

WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS

Phase shift parameter ρ Phase shift parameter ρ

INHERENT OPTICAL PROPERTIES OF NON-SPHERICAL MARINE-LIKE PARTICLES

Phase shift parameter ρ Phase shift parameter ρ

0.5 1.5 1.5

0.5 1 2 5 10 20 50 100 200 γc 0.5 1 2 5 10 20 50 100 200 γa

Optical properties of polydispersions

WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS

Phase shift parameter ρ Phase shift parameter ρ

INHERENT OPTICAL PROPERTIES OF NON-SPHERICAL MARINE-LIKE PARTICLES

Phase shift parameter ρ Phase shift parameter ρ

0.5 1 2 5 10 20 50 100 200 Qb 0.5 1 2 5 10 20 50 100 200 Qbb

WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS

Phase shift parameter ρ Phase shift parameter ρ

0.5 1 2 5 10 20 50 100 200 γb 0.5 1 2 5 10 20 50 100 200 γbb

 0, if D < Dmin or D > Dmax ;

where D0 [µm] is a reference diameter and n0 [ # m −3 µm −1 ] is the differential number

INHERENT OPTICAL PROPERTIES OF NON-SPHERICAL MARINE-LIKE PARTICLES

Phase shift parameter ρ Phase shift parameter ρ

WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS

Phase shift parameter ρ Phase shift parameter ρ

10−3 m = 1.05 + i0.01 2

0.5 1 2 5 10 20 50 200 0.5 1 2 5 10 20 50 200

 0, if D < Dmin or D > Dmax ;

INHERENT OPTICAL PROPERTIES OF NON-SPHERICAL MARINE-LIKE PARTICLES

distribution earlier, to examine how changes in the relative concentration of small to

Results for polydispersions

WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS

INHERENT OPTICAL PROPERTIES OF NON-SPHERICAL MARINE-LIKE PARTICLES

WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS

Phase shift parameter ρ Phase shift parameter ρ

0.6 0.6 Power-law

INHERENT OPTICAL PROPERTIES OF NON-SPHERICAL MARINE-LIKE PARTICLES

Observations on the effects of particle shape on IOPs

WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS

Figure 16 Measurements of near-forward scattering. A is a comparison of the shape of the near-forward

Summary and future prospect

INHERENT OPTICAL PROPERTIES OF NON-SPHERICAL MARINE-LIKE PARTICLES

WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS

INHERENT OPTICAL PROPERTIES OF NON-SPHERICAL MARINE-LIKE PARTICLES

WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS

INHERENT OPTICAL PROPERTIES OF NON-SPHERICAL MARINE-LIKE PARTICLES

WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS

INHERENT OPTICAL PROPERTIES OF NON-SPHERICAL MARINE-LIKE PARTICLES

Symbol Deﬁnition Dimension

WILHELMINA R. CLAVANO, EMMANUEL BOSS & LEE KARP-BOSS

Symbol Deﬁnition Dimension

Oceanography and Marine Biology: An Annual Review, 2007, 45, 39-88

GLOBAL ECOLOGY OF THE GIANT KELP MACROCYSTIS:

Moss Landing, California 95039, U.S.

Universidad Católica del Norte, Coquimbo, Chile

Universidad de Los Lagos, Casilla 557, Puerto Montt, Chile

MICHAEL H. GRAHAM, JULIO A. VÁSQUEZ & ALEJANDRO H. BUSCHMANN

Chile Tristan de Cunha Is. South Australia New Zealand

MICHAEL H. GRAHAM, JULIO A. VÁSQUEZ & ALEJANDRO H. BUSCHMANN

Organismal biology of Macrocystis