Journal of Biogeography (J. Biogeogr.
) (2004) 31, 1107–1124
ORIGINAL
ARTICLE
Charles Darwin Research Station, Santa Cruz,
Galapagos, Ecuador
*Correspondence: G. J. Edgar, School of
Zoology, Tasmanian Aquaculture and Fisheries
Institute, University of Tasmania, GPO Box 252-
05, Hobart, Tasmania 7001, Australia.
E-mail: g.edgar@utas.edu.au
Present address: J. M. Fariña, Center for
Advanced Studies in Ecology and Biodiversity
(CASEB), Pontificia Universidad Católica de
Chile, Santiago, Chile and Department of
Ecology and Evolutionary Biology, Brown
University, Providence, RI 02912, USA.
ª 2004 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi
Regional biogeography of shallow reef
fish and macro-invertebrate communities
in the Galapagos archipelago
G. J. Edgar*, S. Banks, J. M. Fariña , M. Calvopiña and C. Martı́nez
ABSTRACT
Aim To delineate biogeographical patterns in Galapagos shallow-water reef
fauna at regional scales.
Location Galapagos Islands.
Methods Fishes and macro-invertebrates were quantitatively censused using
underwater visual techniques along more than 500 transects at defined depth
strata across the Galapagos archipelago. Data were analysed using multivariate
techniques to define regional patterns and identify species typical of different
regions.
Results Subtidal communities of fishes and macro-invertebrates on shallow
reefs differed consistently in species composition across the Galapagos
archipelago, with three major biogeographical groupings: (1) the ‘far-northern
area’ containing the islands of Darwin and Wolf, (2) the ‘central/south-eastern
area’, including the east coast of Isabela, and (3) the ‘western area’, encompassing
Fernandina and western Isabela. In addition, the northern islands of Pinta,
Marchena and Genovesa form a separate region in the central/south-eastern area,
and Bahia Elizabeth and Canal Bolivar separate from other parts of the western
area. The far-northern bioregion is characterized by high fish species richness
overall, including a high proportion of species of Indo-Pacific origin. However,
very few endemic fishes or species with distributions extending south from
Ecuador (‘Peruvian’ species) are present, and the bioregion also possesses
relatively low species richness of mobile macro-invertebrate taxa. By contrast, the
‘western’ bioregion possesses disproportionately high numbers of endemic fish
taxa, high numbers of cool-temperate Peruvian fish species, and high invertebrate
species richness, but very few species of Indo-Pacific origin. The Bahia Elizabeth/
Canal Bolivar bioregion possesses more endemic species and fewer species with
Peruvian affinities than coasts within the western bioregion. The northern
bioregion of Pinta, Marchena and Genovesa represents an overlap zone with
affinities to both the far-northern and south-eastern islands. The south-eastern
bioregion includes species from a variety of different sources, particularly
‘Panamic’ species with distributions extending north to Central America.
Main conclusions On the basis of congruent divisions for reef fish and macro-
invertebrate communities, the Galapagos archipelago can be separated into three
major biogeographical areas, two of which can be further subdivided into two
regions. Each of these five bioregions possesses communities characterized by a
distinctive mix of species derived from Indo-Pacific, Panamic, Peruvian and
endemic source areas. The conservation significance of different regions is not
reflected in counts of total species richness. The regions with the lowest overall
fish species richness possess a temperate rather than tropical climate and highest
levels of endemism.
1107
G. J. Edgar et al.
Keywords
Benthos, Galapagos, fishes, macro-invertebrates, reef, biogeographical region,
species richness, endemism, faunal distribution.
INTRODUCTION
The more than 100 volcanic islands and islets of Galapagos
possess one of the most interesting biogeographical settings on
earth. Galapagos is the only tropical archipelago lying at the
intersection of major warm- and cool-water current systems,
being located in the path of (1) the warm south-westerly
flowing Panama Current, (2) the cool north-westerly flowing
Peru current, and (3) the cold eastward-flowing subsurface
equatorial undercurrent (Houvenaghel, 1978; Banks, 2002).
The latter rises to the surface along western and southern
margins of the Galapagos Plateau, generating highly produc-
tive upwelling systems (Pak & Zanfield, 1974). As a
consequence, oceanographical conditions vary markedly over
short spatial scales across the archipelago, with quite extreme
environmental differences existing between the cool upwelling
region in the west and the warm oligotrophic north. For
example, water temperature at 10 m depth at Punta Espinosa
on the westernmost island of Fernandina averaged 18 C
compared with 24 C 180 km north off the island of Wolf
between June 1996 and February 1997 (Wellington et al.,
2001). Much of this regional variation in ocean climate
disappears during El Niño years (Banks, 2002), when water
temperatures rise above 25 C throughout the archipelago for
periods that can exceed a year (Wellington et al., 2001).
The anomalous ocean climate surrounding Galapagos
translates to distinctive marine ecosystems that occur in close
proximity, and a biota that includes elements characteristic of
tropical (e.g. manta rays, reef sharks, corals), temperate (sea
lions, kelp) and even subantarctic (fur seals, penguins,
albatross) seas. A large component of endemic species is
present (Bustamante et al., 2000), notably including the marine
iguana (Amblyrhynchus cristatus), flightless cormorant (Phala-
crocorax harrisi) and the only tropical laminarian kelp (Eisenia
galapagensis).
Despite this scientific importance, little has been published
on the distribution of marine biodiversity across the region
(Bensted-Smith, 2002). The most widely quoted marine
regionalization for the archipelago was described more than
30 years ago by Harris (1969). This regionalization was
originally physical rather than biological, with water tempera-
ture data used as a surrogate for biological data because the
latter were largely lacking (Harris, 1969). In the Harris scheme,
Galapagos waters were subdivided into five regions (north,
west, south, central and central mixing). Jennings et al. (1994)
provided some support for Harris’ (1969) scheme with
information on reef fishes; however, only 10 sites were
investigated during that study and insufficient data were
available to compare variation within as well as between
regions. In the sole major archipelago-wide investigation prior
1108 Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd
to the present study, Wellington (1975) considered that four
major biogeographical regions were clearly defined for Galap-
agos (western, southern, central and northern).
The present study aimed to clarify broad-scale marine
biogeographical patterns across Galapagos. It was initiated
partly for scientific interest and partly in response to a
management need. All waters extending 40 miles offshore from
an imaginary line joining the outer islands are regulated by the
Ecuadorian Government for conservation of biodiversity
within the Galapagos Marine Reserve (GMR). The GMR is
regulated by a Management Plan, which states as a principal
aim that biodiversity will be protected in all biogeographical
regions. The Management Plan was negotiated by local
stakeholders, who recognized that marine plants and animals
required protection within ‘no-take’ sanctuary zones of
adequate size in all different bioregions. This aim could only
be achieved if biogeographical regions were more accurately
defined than currently in the Harris scheme.
METHODS
Sites surveyed
Quantitative data were collected between 13 May 2000 and
13 December 2001 during research cruises to define baseline
conditions in different management zones within the GMR.
The shallow subtidal rocky reef habitat investigated here, and
used as the data set for deriving the regionalization, represents
the predominant habitat type around the Galapagos coastline
(> 95% of shallow habitat, Bustamante et al., 2002). Shallow
reefs also comprise the habitat most affected by fishing and
other human activity.
Underwater visual censuses along line transects were under-
taken during daylight hours at 50 islands and islets distributed
across the archipelago (Fig. 1). Generally, two different depth
contours were surveyed at a single site. For some sites, the two
depth strata surveyed were parallel and immediately adjacent to
each other, while in other areas depth strata were offset by up to
300 m when divers were working from different boats.
Consequently, the term ‘site’ is somewhat ambiguous, and
the term ‘depth strata’ preferred, referring to one depth interval
at a site. Overall, a total of 579 and 569 depth strata were
surveyed for fish and macro-invertebrates, respectively.
Faunal survey protocols
Fish surveys were undertaken by laying a 50 m transect line
along a defined depth contour within the range from 2 to 20 m
depth. A diver swam beside the transect line at a distance of
2.5 m, recording on a waterproof notepad the abundance of
 Regional biogeography of Galapagos shallow-water reef fauna
Figure 1 Sites with quantitative fish and
invertebrate transects surveyed during 2000
and 2001. The majority of sites are overlap-
ping and not shown. Island names are in bold
italics, site names in regular font.
fishes in a 5 m wide swathe (i.e. from the transect line out to a
distance of 5 m). This process was then repeated on return
along the other side of the transect line, with data from the two
adjoining sides of the transect added together for each
50 m · 10 m census block. For the majority (59%) of depth
strata, two replicate 50 m · 10 m blocks were surveyed and
mean data for that depth stratum used in analyses; however, on
the remaining occasions the census block was not duplicated.
Macro-invertebrates were censused along the same transect
lines as for fishes. However, only a 1 m wide swathe up and
back along the transect line was surveyed by the diver, who
recorded the abundance of macro-invertebrate species (sea
stars, sea cucumbers, sea urchins, octopus, large gastropods,
large bivalves, spiny lobsters and large crabs) in each
2 m · 50 m block. As with analyses of fishes, the majority of
depth strata were duplicated, and the mean value for the two
blocks used in analyses.
Analyses
The number of species recorded during 50 m transects at
depth strata across the archipelago was plotted in Arcview
(ESRI, Redlands, CA, USA). Smoothed contour plots were
produced using the Inverse Distance Weighted interpolation
function with 24 nearest neighbours along the coastline.
Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd
Offshore extrapolations have been masked in figures using a
10 km corridor off the coast.
Faunal relationships between sites were investigated using
non-metric multidimensional scaling (MDS) plots calculated
by PRIMER (Plymouth, UK; Carr, 1996). These provided the
best graphical depictions in two dimensions of biological
similarities between sites.
Because data for individual transects were greatly affected by
local environmental conditions, particularly depth and wave
exposure, and the aim of the study was to identify regional
patterns, site data were grouped by calculating a mean value
for each 10 depth strata in nearest proximity around the coast
of each island. Data matrices showing mean abundances of
different fish or invertebrate species at different sites used for
MDS plots were initially fourth root-transformed, then
converted to matrices of biotic similarity between pairs of
sites using the Bray–Curtis similarity index, as recommended
by Faith et al. (1987). The usefulness of the two-dimensional
MDS display of biotic relationships is indicated by the stress
statistic, which signifies a good depiction of relationships when
< 0.1 and poor depiction when > 0.2 (Clarke, 1993).
Additionally, the SIMPER module of PRIMER was used to
identify species that contributed most substantially to the
average similarity within each biogeographical region, and
thereby typified each region (Clarke, 1993).
1109
G. J. Edgar et al.
Faunal patterns across the archipelago were also analysed
using canonical analysis of principal coordinates (CAP)
(Anderson, 2003), a constrained ordination procedure that
initially calculates unconstrained principal coordinate axes,
followed by canonical discriminant analysis on the principal
coordinates to maximize separation between predefined
groups (Anderson & Robinson, 2003; Anderson & Willis,
2003). CAP is considered more flexible than direct canonical
discriminant analysis because any dissimilarity measure can be
utilized rather than only Mahalanobis distance. Data were
fourth root-transformed and the Bray–Curtis similarity index
used. In order to reduce local effects of depth, data input into
the CAP analysis comprised mean density values of different
species for each 0.01 latitude · 0.01 longitude grid cell
(¼ 1 km2 area). Data were available for 220 grid cells for fish
and 191 grid cells for macro-invertebrates.
The CAP procedure was also used to calculate miscalcula-
tion error for bioregional groupings using the leave one-out
approach. Each grid cell point was removed from the data set,
the CAP analysis rerun using the remaining observations, and
then the removed data point classified to the nearest group
centroid in canonical space. Comparison of known with
allocated groups provided miscalculation error (Anderson &
Willis, 2003).
RESULTS
A total of 175 fish and 47 macro-invertebrate species were
recorded during quantitative surveys. Assemblages of fishes
sighted in the various transect blocks exhibited consistent
regional patterns. This is indicated in the MDS plot of Fig. 2,
where data for each point represent a mean value calculated
from 10 adjacent depth strata values, and distance between
points reflects faunal similarity.
Figure 2 MDS plot showing relationships between fish faunas
censused in different islands of Galapagos. Each point comprises
the mean of 10 depth strata, and is coded by island prefix and in
sequential order around each island. Stress ¼ 0.16.
1110 Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd
Figure 3 MDS plot showing relationships between fish faunas in
different islands of Galapagos. This analysis is the same as for
Fig. 2, except that data for Isabela (solid line) and Fernandina
(dotted line) were grouped at intervals of three rather than 10
depth strata. Adjoining locations on Isabela and Fernandina are
connected by line. Stress ¼ 0.16. Arrows indicate prominent site
locations for the two islands.
Fish assemblages observed off the island of Darwin and in
three different areas off Wolf lie almost on top of each other in
the graph, indicating that they possess almost identical fish
faunas. This ‘far-northern’ grouping is closest to the fish fauna
found off Pinta, Marchena and Genovesa. By contrast, data for
sites off Fernandina and Isabela lie at the opposite extreme
from Darwin and Wolf on the graph. The fish fauna off
western Isabela shows virtually no similarity to that found in
the far-northern region. The separation between individual
sites on Fernandina and Isabela is very wide, whereas the
faunas on other islands are much more homogeneous.
In order to more clearly classify the huge variability in the
fish fauna that occurred around Fernandina and Isabela, the
data set was re-analysed with finer subdivision of sites for these
two islands. Instead of calculating an average for every
10 depth strata around the coast, an average was calculated
for every three depth strata. Sites adjoining each other around
the coastlines of Fernandina and Isabela have been connected
by line in Fig. 3.
The fish fauna of Isabela is seen in Fig. 3 to possess three main
components: (1) an east coast fauna that is most distinctive
between Cabo Marshall and Bahia Cartago, and which is similar
to the faunas of Santiago, Santa Cruz, Floreana and Pinzon, (2)
an exposed coast fauna that is most distinctive at Punta Vicente
Roca and to a lesser extent Caleta Iguana, and which overlaps
with the west coast fauna of Fernandina, and (3) a sheltered west
coast fauna that is most distinctive in Bahia Elizabeth near Islas
Marielas, and which is similar to the fish fauna on the east coast
of Fernandina between Punta Mangle and Punta Espinosa. The
eastern Isabela fauna appears to separate from the western fauna
at a site just west of Punta Albermarle in the north and just east
of Isla Tortuga in the south.
 Regional biogeography of Galapagos shallow-water reef fauna
Figure 4 MDS plot showing relationships between macro-
invertebrate faunas censused in different islands of Galapagos.
Each point comprises the mean of 10 depth strata, and is coded by
island prefix and in sequential order around each island.
Stress ¼ 0.15.
Analysis of the macro-invertebrate data set (Fig. 4) reveals
three major groups of sites: (1) Darwin and Wolf, (2)
Fernandina and western Isabela, and (3) other islands.
Genovesa and Pinta also separate from the main island
grouping because of their greater affinity with the fauna of
Darwin and Wolf. The fauna of Marchena is quite variable but
with a high level of similarity to central and southern islands
and north-eastern Isabela. The fauna of Pinzon is distinctive.
The macro-invertebrate fauna also shows a very high degree
of variation around Isabela, but is more homogeneous around
Fernandina than was seen for the fish fauna (Fig. 5). The
invertebrate fauna off the coast of Isabela from Punta
Albermarle to Cuatro Hermanos overlaps the fauna of
Floreana, Santiago, Santa Fe, Santa Cruz, San Cristobal,
Figure 5 MDS plot showing relationships between macro-
invertebrate faunas in different islands of Galapagos. This analysis
is the same as for Fig. 4, except that data for Isabela (solid line)
and Fernandina (dotted line) were grouped at intervals of three
rather than 10 depth strata. Adjoining sites on Isabela and Fer-
nandina are connected by line. Stress ¼ 0.15. Arrows indicate
prominent site locations for the two islands.
Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd
Espanola and Rabida, while the west coast fauna exhibits
major differences between Bahia Elizabeth (Islas Marielas) and
Caleta Iguana.
Patterns of fish species richness (i.e. the number of fish
species observed per 50 m · 10 m transect block) across the
archipelago generally reflected clinal trends in communities
identified by MDS. Fish species richness was highest around
the far-northern islands of Darwin and Wolf and lowest off
Fernandina, Santa Cruz, and the Bahia Elizabeth region of
western Isabela (Fig. 6). However, the distribution of endemic
Galapagos fishes was opposite to that seen for the total fauna,
with endemic species present in highest numbers near Islas
Marielas in Bahia Elizabeth and also off western Isabela,
Fernandina, Santa Fé and south-western Floreana (Fig. 7).
The unusually high species richness in the far-northern
region was caused by the presence in that area of numerous
species with ranges extending westward across the Indo-Pacific
(Fig. 8a). Many of these species were coral reef-associated
wrasses, butterflyfishes, pufferfishes and jacks. The far-nor-
thern islands also possessed a disproportionately high number
of species with ‘Panamic’ ranges that extend north of Ecuador
but not south (Fig. 8b); nevertheless, in addition to the virtual
absence of endemic Galapagos fishes (Fig. 7), this region
included very few species with ‘Peruvian’ ranges extending
south along the South American coast (Fig. 8d).
Southward-ranging Peruvian species were largely restricted
to western, northern and southern Fernandina, and south-
western and north-western Isabela (Fig. 8d), probably because
most of these species associate with seaweed habitats that are
largely absent elsewhere in the archipelago. Fish species with
wide South American ranges that extended both north and
south of Ecuador were widespread throughout the archipelago
(Fig. 8c), except for disproportionately low numbers off Bahia
Elizabeth, Fernandina (particularly the east coast) and Santa
Cruz.
In contrast to the situation with fishes, the number of
macro-invertebrate species recorded per 50 m · 2 m transect
block varied relatively little across the archipelago, with a
general average of 6.0 species per transect (Fig. 9). Neverthe-
less, 25% fewer species occurred around the far-northern
islands of Darwin and Wolf (4.5 species per transect) than
elsewhere.
Invertebrate species with Indo-Pacific distributions occurred
throughout the archipelago, with lowest species richness in the
south-west (Fig. 10a). Species with ranges only to the north of
Ecuador were also widely distributed but with no decline
apparent in the western region (Fig. 10b). The relatively low
species richness of macro-invertebrates around the two
northernmost islands was caused by few species with wide-
spread South American distributions occurring in the far-
northern area (Fig. 10c). Widespread South American
species were, however, disproportionately abundant off western
Isabela and Fernandina, and presumably tolerated cooler
conditions than the other regional species groups.
No macro-invertebrate species were sighted that possessed a
distribution on the South American continent solely south
1111
G. J. Edgar et al.

92° 91° 90° 89°

2° 2° Management zones
Conservation
Tourism
Fishing
Port

1° 1°
Species
10
11
12
13
14
0° 0° 15
16
17
18
19
20
21
22 Figure 6 Shaded contour plot showing
1° 1° 23
mean total number of fish species observed
per 50 m transect around different coasts of
Galapagos. Line width of island outline
reflects management zone of that section of
92° 91° 90° 89° coast.

92° 91° 90° 89°

2° 2° Management zones
Conservation
Tourism
Fishing
Port

1° 1°
Species
0
0.2
0.4
0.6
0.8
0° 0° 1.0
1.2
1.4
1.6

1° 1°

Figure 7 Shaded contour plot showing

mean total number of endemic Galapagos
92° 91° 90° 89° fish species per 50 m transect.

from Ecuador. Moreover, only four endemic invertebrate
species were recorded in transects – the slipper lobster
Scyllarides astori, the sea urchin Eucidaris galapagensis, the
octopus Octopus oculifer and the scallop Nodipecten magnificus,
hence plots of macro-invertebrates comparable with Figs 7 and
8d could not be depicted.
Generally, species richness analyses for fish and invertebrate
data sets were consistent with MDS analyses in showing that
Galapagos coastal waters are best divided into the five marine
biogeographical regions (bioregions) shown in Fig. 11. These
bioregions are referred to as ‘far-northern’, ‘northern’, south-
eastern’, ‘western’ and ‘Elizabeth’. The name ‘Elizabeth’ has
been used because the core features of that bioregion are most
evident within Bahia Elizabeth rather than Canal Bolivar,
which possess more overlap with the ‘western’ bioregion. The
Elizabeth bioregion extends from Punta Espinosa and the
northern point of Tagus Cove in the north to Punta Mangle
and just east of Punta Moreno in the south. The western
bioregion on Isabela extends from just west of Punta
Albermarle to just east of Isla Tortuga.
The mean densities of common fish and invertebrate species
in different bioregions are listed in Tables 1 and 2. Species
disproportionately abundant or rare in each bioregion, as
identified using SIMPER analysis (Clarke, 1993), are also

1112 Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd

 Regional biogeography of Galapagos shallow-water reef fauna

92° 91° 90° 89° 92° 91° 90° 89°

2° 2° 2° 2°

1° 1° 1° 1°

0° 0° 0° 0°

1° 1° 1° 1°

(a) (b)
92° 91° 90° 89° 92° 91° 90° 89°
92° 91° 90° 89° 92° 91° 90° 89°

2° 2°

1° 1°

0° 0°

1° 1°

(c) (d)

92° 91° 90° 89° 92° 91° 90° 89°

Figure 8 Shaded contour plot showing mean total number of fish species per 50 m transect for species with Panamic (ranges extending
northwards along the South American coast but not southwards from Ecuador), Indo-Pacific (ranges extending westwards to at least
Hawaii), Peruvian (ranges extending southwards along the South American coast but not northwards from Ecuador) and widespread
(ranges extending both northwards and southwards along the South American coast from Ecuador) ranges.

indicated. Note that species showing disproportionately high
mean abundances within a bioregion are not necessarily
identified by SIMPER analysis as typifying that bioregion. For
example, the endemic goby Lythrypnus gilberti is most
abundant in the south-eastern bioregion but better typifies
the Elizabeth bioregion. This fish occurred in extremely high
abundance (> 1 m)2) at a few sites on south-western Isabela in
the south-eastern bioregion but was absent from the majority
of sites, whereas the species occurred in lower maximal
abundance in the Elizabeth bioregion but at a high proportion
of sites.
Faunal abundance data for different biogeographical regions
exhibited similar patterns to species richness data in that
densities of Indo-Pacific species were generally highest in the
far-northern bioregion, endemic species were most abundant
in the Elizabeth and western bioregions, and Peruvian species
were most abundant in the western bioregion. Nevertheless, a
few anomalous distributions existed. For example, the Indo-
Pacific pufferfish Sphoeroides annulatus was not recorded in
the far-northern islands while the parrotfish Scarus ghobban
was widely distributed throughout the archipelago rather than
being concentrated in the far north. In addition, although the
echinoid Echinometra vanbrunti and the holothurian Holothu-
ria difficilis possess Panamic distributions, these species were
not detected in the warm far-northern region of Galapagos.
Faunal data pertaining to different 0.01 latitude and
longitude grid cells have been classified in multidimensional
space using CAP analysis to maximize differences between the
five bioregional groups. The best congruence with the five
bioregional groups was found using the first 20 principal

Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd 1113

G. J. Edgar et al.
92° 91° 90° 89°
2°
1°
0°
1°
92° 91° 90° 89°
components for fishes and first 10 principal components for
macro-invertebrates. Differences between bioregional groups
were statistically significant for all pairwise comparisons
(permutation test, P < 0.001, including Bonferoni correction
for 10 pairwise tests).
For both fishes (Fig. 12) and macro-invertebrates (Fig. 13),
the first two canonical axes did not separate the western and
Elizabeth regions (Figs 12a and 13a); hence the third canonical
axes have been included in figures (Figs 12b and 13b).
Although these figures indicate a stronger separation between
the western and Elizabeth bioregions for fishes than inverte-
brates, the leave-one-out procedure indicates a stronger
separation between these bioregions for the macro-invertebrate
data when all canonical axes are considered (Tables 3 and 4).
Data for each grid cell were accurately classified into the
appropriate bioregional group in the majority of cases using
the leave-one-out procedure, with misclassification primarily
occurring between the western and Elizabeth bioregions, and
between the northern and south-eastern bioregions. Thus, a
primary three-group separation exists between far-northern,
northern + south-eastern, and western + Elizabeth bioregions.
Species showing high correlations with the major canonical
axes included nearly all species identified by SIMPER analysis
as typifying particular regions. A large group of species,
including the moray eel Gymnothorax dovii, surgeonfishes
Acanthurus nigricans and Prionurus laticlavius, priacanthid
Heteropriacanthus cruentatus, trumpetfish Aulostomus chinen-
sis, Moorish idol Zanclus cornutus, pufferfishes Arothron
meleagris and Canthigaster punctatissima, wrasse Thalassoma
lucasanum, triggerfish Sufflamen verres, sea urchin Diadema
mexicanum, spiny lobster Panulirus penicillatus and crab
Percnon gibbesi, were principally confined to the far-northern
islands. The majority of these species possess Indo-Pacific
distributions.
1114
2°
1°
0°
1°
Figure 9 Shaded contour plot showing
mean total number of invertebrate species
observed per 50 m transect around different
coasts of Galapagos.
The groupers Paralabrax albomaculatus and Mycteroperca
olfax, goby Lythrypnus gilberti, sea cucumber Stichopus fuscus,
and sea stars Nidorellia armata and Pharia pyramidata were
most highly correlated with the Elizabeth bioregion; the wrasse
Halichoeres dispilus, weedfish Labrisomus dendriticus, and sea
urchins Lytechinus semituberculatus and Centrostephanus cor-
onatus with the western bioregion; and the wrasses Semicossy-
phus darwini and Bodianus eclancheri, hornshark Heterodontus
quoyi, and knifejaw Oplegnathus insignis were disproportion-
ately abundant in both these bioregions. Species highly
correlated with the south-eastern region included the dam-
selfish Abudefduf troschelii, wrasse Halichoeres nicholsi, grunt
Haemulon scudderi, and sea urchin Tripneustes depressus. The
holothurians Holothuria atra and Holothuria fuscocinerea were
primarily associated with the northern bioregion.
DISCUSSION
Biogeographical patterns
Faunal abundance and species richness data both indicate that
Galapagos inshore reef ecosystems are divisible into three
major regional components: (1) the far-northern islands of
Darwin and Wolf, (2) the central, southern and eastern islands,
including the east coast of Isabela, and (3) Fernandina and
western Isabela. This basic subdivision was evident in all
analyses. The far-northern area included a large component of
Indo-Pacific and Panamic species but virtually no endemic
Galapagos animals, the western area included many endemic
species and those ranging south down the South American
coast, while the central/eastern/southern area included a
mixture of species derived from various source regions.
In addition, for the central/southern/eastern region, the
majority of analyses indicated that the northern islands of
Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd
 Regional biogeography of Galapagos shallow-water reef fauna

(a) (b)

(c)

Figure 10 Shaded contour plot showing mean total number of invertebrate species per 50 m transect for species with Panamic (ranges
extending northwards along the South American coast but not southwards from Ecuador), Indo-Pacific (ranges extending westwards to at
least Hawaii) and widespread (ranges extending both northwards and southwards along the South American coast from Ecuador) ranges.

Pinta, Marchena and Genovesa possessed a fauna distinctively
different from that found on the other islands. These three
northern islands comprised an overlap zone, and had dispro-
portionately more animals of Indo-Pacific origin than islands
further south, but considerably less than in the far-northern
region. Very few endemic Galapagos species or those with a
Peruvian distribution occurred there.
Moreover, in the western area, the region from Bahia
Elizabeth to Punta Espinosa possessed an anomalously high
proportion of endemic fish species, while north-western and
south-western Isabela and western Fernandina possessed a
different fish assemblage type (see Fig. 3) with an unusually
large component of species with Peruvian affinities. For this
reason, and because a number of endemic invertebrate species
have been recorded near Tagus Cove but not elsewhere in
Galapagos, such as the coral Tubastraea faulkneri (Humann,
1988) and the seastars Luidia superba and Heliaster solaris
(Hickman, 1998), the ‘Elizabeth’ bioregion has been separated
from the ‘western’ bioregion.
The regional biogeographical scheme outlined here differs in
several features from the scheme proposed by Harris (1969), in
part because our extensive data set has allowed boundaries of
different zones to be defined with greater precision. Whereas
Harris possessed data for few sites and was unable to precisely
define boundaries, the present analysis included data for two
different faunal components (reef fish and invertebrates) from
c. 200 1-km2 grid cells distributed around all main islands. We
note that the pronounced regional difference that separates the
far-northern from the northern islands was not identified by
Harris because he had no information from Darwin or Wolf.
Harris subdivided the central, southern and eastern islands
into three regions, whereas our analysis did not confirm this
separation. Although our survey results indicated differences
between islands – for example, the fish fauna of Española

Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd 1115

G. J. Edgar et al.
Figure 11 The five major marine bioregions identified for Gal-
apagos.
differing from the fish fauna of Santa Cruz and possessing
relatively close affinity to the northern islands (Fig. 2) – these
patterns were not consistent for both the fish and macro-
invertebrate data sets. Furthermore, the variation around
individual islands was considerable (as seen, e.g. in Fig. 4 for
invertebrates around Santa Cruz) and often greater than
variation between islands. Hence, using our fish and inverteb-
rate data sets, we were unable to subdivide the various central,
eastern and southern islands into consistent subgroups.
The distinctiveness of the Elizabeth grouping was an
unexpected outcome of the analysis. In terms of faunistic
affinity, the core of the Elizabeth bioregion around Islas
Marielas differed from Caleta Iguana as much as Caleta Iguana
did from south-eastern islands such as Santa Cruz. Part of the
reason for this distinctiveness can be inferred from satellite
images of the GMR depicting phytoplankton concentration
(Banks, 2002); nearly all images indicate much higher levels of
primary productivity in the Elizabeth bioregion than elsewhere
in the archipelago. This high productivity may give the
ecosystem its distinctive character.
The strong regional divisions in the Galapagos marine
fauna probably reflect both local environmental conditions
and connectivity of larval propagules with external source
regions. Species with Indo-Pacific distributions occur pre-
dominantly in the far-northern region of Galapagos, where
water temperatures are highest, turbidity is low, coral
development is most extensive, and warm currents are likely
to first strike the archipelago. Many amongst the diverse far-
northern component of species probably maintain gene flow
across the East Pacific Barrier, particularly during El Niño
years when currents from the north-east and north prevail
(Glynn & Ault, 2000). In addition to long-lasting larval
stages, Indo-Pacific fishes in Galapagos often penetrate much
1116
more widely through the archipelago during periods of El
Niño, and possess small local population sizes and distorted
size structures (Grove, 1985). Some species (most notably the
wrasse Stethojulis bandanensis) apparently recruit to the
archipelago episodically only during El Niño years (Victor
et al., 2001).
Populations of Panamic species are probably largely self-
sustaining in Galapagos but with intermittent recruitment
from the mainland and islands to the north. Amongst the local
fish species of Panamic origin where the duration of larval
stage has been examined, some individuals have been found to
settle onto reefs after periods insufficient for long-distance
transport, whereas others possess long larval stages and
predominantly recruit during El Niño years (Wellington &
Victor, 1992; Meekan et al., 2001).
The Peruvian component of species is probably largely self-
sustaining within Galapagos rather than reliant on larval
propagules arriving from the continent. Populations of Peru-
vian species primarily occur in western rather than south-
eastern Galapagos – the geographically closer location to the
southern South American coast. Peruvian species presumably
prefer the west because environmental conditions are colder
with high primary production, and thus more similar to
prevailing environmental conditions in central South America.
All of the endemic species encountered during surveys are
closely related to species present on the South American coast
(see e.g. Lessios et al., 1999).
Patterns of species richness across the archipelago were
dissimilar for fishes and macro-invertebrate, in part because
macro-invertebrate species with Peruvian affinities were
absent and endemic invertebrate species were depauperate.
Representation of these groups within the Galapagos fauna
has probably declined greatly during recent decades, partic-
ularly following the 1982/83 El Niño. This is indicated, for
example, by a collapse in populations of the endemic scallop
Nodipecten magnificus, a large mollusc that occurred com-
monly in the Elizabeth bioregion until 1983. In that year, all
living individuals monitored by researchers died (Robinson,
1985). Scallop numbers have shown little subsequent recov-
ery, with only three individuals recorded during recent
surveys. Several other endemic invertebrate species, including
the corals Tubastraea floreana Wells and Madrepora galap-
agensis Vaughan and the seastar Heliaster solaris (A.H. Clark),
have not been sighted since 1983 (Cairns, 1991; Hickman,
1998).
The 1982/83 El Niño clearly had a catastrophic impact on
Galapagos marine biodiversity, with consequences that persist
today (Bensted-Smith, 2002). Macroalgae and invertebrates
with cool-temperate affinities appear to have been most
affected (also corals: Glynn, 1994), their ranges contracting
greatly and populations now being largely confined to localized
areas in the west. Thus, patterns of invertebrate biodiversity
identified would perhaps have been quite different if our
surveys had been conducted in 1980.
The poor recovery of marine invertebrates following El Niño
compared with fishes is likely due to limited metapopulation
Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd
 Regional biogeography of Galapagos shallow-water reef fauna

Table 1 Mean abundance per 500 m2 transect in different biogeographical regions of fish species recorded on at least five transects.
Fish species identified using SIMPER analysis to be associated with particular bioregions are listed by superscript in rank order of strength of
association, with negative numbers indicating negative association

Biogeographical region

Species Far-northern Northern South-eastern Western Elizabeth Overall

Endemic species
Lythrypnus gilberti (Heller & Snodgrass) 0 0 24.6 1.40 17.62 15.2
Xenocys jessiae Jordan & Bollman 0 0.63 6.87 41.0 20.3 12.1
Girella freminvillei (Valenciennes) 0.02 0.60 2.64 2.57 2.01 2.09
Orthopristis forbesi Jordan & Starks 0 0.10 4.38 1.62 1.88 2.80
Lepidonectes corallicola (Kendall & Radcliffe) 0.16 0.26 1.47 4.45 2.514 1.77
Acanthemblemaria castroi Stephens & Hobson 0 0 2.20 1.56 0.01 1.41
Mugil rammelsbergi (Ebeling) 0 0.10 0.39 0 8.06 1.06
Paralabrax albomaculatus (Jenyns) 0 0.46 0.15 0.09 1.345 0.30
Sphoeroides angusticeps (Jenyns) 0 0.10 0.10 0.02 0.07 0.08
Odontoscion eurymesops (Heller & Snodgrass) 0 0 0 0.35 0 0.05

Panamic species
Thalassoma lucasanum (Gill) 1027.9 158.8 44.7)1 28.0)3 4.67)3 123.2
Prionurus laticlavius Valenciennes 123.2 141.5 160.3 24.6)2 6.58)2 118.6
Stegastes beebei (Nichols) 21.6)2 76.5 75.8 118.0 107.7 81.8
Apogon atradorsatus Heller & Snodgrass 0.08 0.08 44.0 2.35)10 3.32)10 24.3
Stegastes arcifrons (Heller & Snodgrass) 13.9 22.5 7.28 0.21)11 2.51 8.30
Microspathodon dorsalis (Gill) 0.91 6.61 7.94 7.79 0.22 6.43
Johnrandallia nigrirostris (Gill) 7.41 7.93 7.98 0.67)5 0.32)7 6.03
Haemulon scudderi Gill 0 0 10.5 1.93 1.04 6.00
Mycteroperca olfax (Jenyns) 0.06 1.74 2.20 13.6 4.28 3.92
Lutjanus viridis (Valenciennes) 0.33 1.96 5.82 1 0.01 3.56
Halichoeres nicholsi (Jordan & Gilbert) 0.07 0.69 4.17 1.16)8 0.56)9 2.56
Labrisomus dendriticus (Reid) 0.60 1.22 1.22 9.60 2.30 2.55
Cirrhitus rivulatus Valenciennes 1.41 1.96 1.50 3.03 1.33 1.77
Chromis alta Greenfield & Woods 0.19 0.43 2.28 1.06 0.35 1.49
Lutjanus aratus (Günther) 0 0.63 1.86 0 0 1.08
Scarus compressus (Osborn & Nichols) 0.18 1.31 1.11 0.66 0.66 0.96
Microspathodon bairdii (Gill) 0.95 0.58 0.79 0.82 2.33 0.94
Thalassoma grammaticum Gilbert 1.95 0.20 0.07 2.58 0 0.59
Kyphosus elegans (Peters) 1.50 0.44 0.27 0.29 0.06 0.36
Cephalopholis panamensis (Steindachner) 0.36 0.41 0.41 0.09 0.03 0.32
Dermatolepis dermatolepis (Boulenger) 1.65 0.25 0.23 0.16 0.01 0.30
Canthigaster punctatissima (Günther) 0.74 0.71 0.23 0 0 0.28
Gymnothorax dovii (Günther) 1.90 0.82 0.04 0 0 0.27
Sargocentron suborbitalis (Gill) 0.05 0.25 0.05 0.53 0.85 0.24
Euthynnus lineatus Kishinouye 0 1.06 0.12 0.07 0 0.22
Myripristis leiognathos Valenciennes 0.03 0.39 0.31 0 0 0.22
Lutjanus novemfasciatus(Gill) 0.17 0.18 0.2 0 0 0.14
Dasyatis brevis (Garman) 0 0.02 0.14 0.08 0.03 0.09
Coryphopterus urospilus Ginsburg 0 0.02 0.15 0.07 0 0.09
Synodus lacertinus Gilbert 0.17 0.01 0.08 0.01 0.12 0.07
Scorpaena mystes Jordan & Starks 0.15 0.04 0.06 0.06 0 0.06
Hypsoblennius brevipinnis (Günther) 0 0 0.08 0.07 0 0.05
Hoplopagrus guentheri Gill 0.07 0.01 0.01 0 0 0.01

Peruvian species
Anisotremus scapularis (Tschudi) 0.12 0.02 1.21 17.15 5.52 3.81
Bodianus eclancheri (Valenciennes) 0 0 0.26 9.486 0.74 1.65
Sphyraena idiastes Heller & Snodgrass 0 0 1.72 0.33 0.60 1.03
Oplegnathus insignis (Kner) 0.10 0 0.14 4.01 0.81 0.77
Semicossyphus darwini (Jenyns) 0 0 0.03 2.84 0.49 0.49
Heterodontus quoyi (Freminville) 0 0 0 0.12 0.15 0.03

Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd 1117

G. J. Edgar et al.

Table 1 continued

Biogeographical region

Species Far-northern Northern South-eastern Western Elizabeth Overall

Indo-Pacific species
Scarus ghobban Forsskal 0.58 8.12 5.20 1.84)7 2.29 4.47
Cirrhitichthys oxycephalus (Bleeker) 5.616 1.22 1.51 0.77 0.08 1.50
Acanthurus nigricans Linnaeus 19.03 0.47 0.17 0 0 1.49
Sphoeroides annulatus (Jenyns) 0 0.01 2.47 0.16 0.26 1.37
Zanclus cornutus (Linnaeus) 2.869 3.806 0.80 0.02 0 1.16
Scarus rubroviolaceus Bleeker 1.24 0.72 0.63 0.02 0.19 0.55
Aulostomus chinensis Lacepede 4.797 0.42 0.08 0 0 0.44
Seriola rivoliana Valenciennes 0.76 1.61 0.27 0 0.01 0.42
Arothron meleagris (Bloch & Schneider) 1.37 1.45 0.16 0.01 0 0.39
Melichthys niger (Bloch) 3.08 0.26 0.17 0 0 0.34
Elagatis bipinnulata (Quor & Gaimard) 0.33 0.40 0.27 0 0 0.23
Fistularia commersonii Rüppell 0.36 0.75 0.10 0.03 0 0.19
Chilomycterus affinis Günther 0 0.01 0.14 0 0.15 0.09
Taeniura meyeri (Müller & Henle) 0.01 0.03 0.10 0.09 0.03 0.07
Sphyrna lewini (Griffith & Smith) 0.79 0.01 0.02 0 0 0.07
Acanthurus xanthopterus Valenciennes 0.02 0.01 0.10 0.02 0.02 0.06
Diodon hystrix Linnaeus 0 0.02 0.07 0.02 0.02 0.05
Heteropriacanthus cruentatus (Lacepede) 0.60 0.08 0 0 0 0.05
Triaenodon obesus (Rüppell) 0.02 0 0.07 0 0 0.04
Muraena lentiginosa Jenyns 0 0.02 0.04 0.03 0.09 0.04
Diodon holocanthus Linnaeus 0 0 0.06 0.04 0.03 0.04
Novaculichthys taeniourus (Lacepede) 0.18 0.04 0.03 0.02 0 0.04
Aetobatus narinari (Euphrasen) 0.05 0 0.02 0.01 0 0.02
Myripristis berndti Jordan & Evermann 0.02 0.02 0.02 0 0 0.02
Ostracion meleagris Shaw 0 0 0.02 0 0 0.01
Bothus leopardinus (Günther) 0.04 0.02 0.01 0 0 0.01
Acanthocybium solandri (Cuvier) 0 0 0.02 0 0 0.01
Carcharhinus galapagensis (Snodgrass & Heller) 0.05 0 0.02 0 0 0.01
Arothron hispidus (Linnaeus) 0 0.02 0 0 0 0.01
Thalassoma purpureum (Forsskal) 0.06 0 0 0 0 0.01
Pseudobalistes naufragium (Jordan & Starks) 0 0 0.01 0.01 0 0.01

Widespread species
Paranthias colonus (Günther) 1033.42 452.6 604.0 234.3)1 164.5)1 511.6
Halichoeres dispilus (Gill) 0.90)1 36.5 38.4)2 90.7 40.26 43.6
Ophioblennius steindachneri Jordan & Evermann 72.84 33.2 20.7 21.0 4.39)4 24.4
Bodianus diplotaenia (Gill) 11.9 15.9 19.3 18.9 21.1)6 18.4
Holacanthus passer Valenciennes 7.77)3 16.0 16.83 7.23)4 2.92)5 13.2
Abudefduf troschelii (Gill) 0.01 2.00)2 21.64 0.83 3.99)8 12.4
Anisotremus interruptus (Gill) 0.1 15.95 16.9 1.30)9 0.05 11.5
Plagiotremus azaleus (Jordan & Evermann) 20.010 6.96)1 9.28 11.9 17.81 11.0
Epinephelus labriformis (Jenyns) 1.91 5.42 6.31 2.77)6 14.1 6.15
Mulloidichthys dentatus (Gill) 0.5 7.45 6.26 8.42 0.51 5.75
Chromis atrilobata Gill 0.03 3.62 6.5 4.52 4.96 5.19
Serranus psittacinus Velenciennes 0.07 5.573 2.61 1.4 1.82 2.58
Trachinotus stilbe (Jordan & Macgregor) 5 0.1 2.11 0 0 1.49
Sufflamen verres (Gilbert & Starks) 4.788 2.744 1.17 0.09 0.28)11 1.39
Orthopristis chalceus (Günther) 0 0.05 2.28 0.34 0.87 1.37
Chaetodon humeralis Günther 0 0.05 1.38 0.27 0.4 0.83
Scarus perrico Jordan & Gilbert 0.08 0.15 1.05 0.58 0.05 0.68
Lutjanus argentiventris (Peters) 0 0.1 0.69 0.27 0.11 0.44
Nicholsina denticulata (Evermann & Radcliffe) 0 0.1 0.08 1.92 0.12 0.36
Kyphosus analogus (Gill) 3.3 0.2 0.05 0.01 0 0.29
Apogon pacificus Herre 0 0 0.21 0 0.81 0.20

1118 Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd

 Regional biogeography of Galapagos shallow-water reef fauna

Table 1 continued

Biogeographical region

Species Far-northern Northern South-eastern Western Elizabeth Overall

Balistes polylepis Steindachner 0.36 0.3 0.2 0.03 0 0.18

Rypticus bicolour Valenciennes 0.05 0.07 0.25 0.06 0 0.16
Malacoctenus tetranemus (Cope) 0.02 0 0.16 0.06 0.05 0.10
Caulolatilus princeps (Jenyns) 0 0 0.02 0.14 0.65 0.10
Scomberomorus sierra Jordan & Starks 0.01 0 0.13 0.02 0.03 0.08
Alphestes immaculatus Breder 0 0.04 0.09 0.02 0.12 0.07
Rypticus nigripinnis Gill 0 0.02 0.11 0 0 0.06
Hippocampus ingens Girard 0 0 0.02 0.04 0.09 0.03
Stegastes acapulcoensis (Fowler) 0 0.01 0.01 0 0.06 0.01

Table 2 Mean abundance per 100 m2 transect in different biogeographical regions of macro-invertebrate species recorded on at least
two transects. Species identified using SIMPER analysis to be associated with particular bioregions are listed by superscript in rank order
of strength of association, with negative numbers indicating negative association. Taxonomic groups are: E, echinoid; A, asteroid;
H, holothurian; C, crustacean; M, mollusc

Biogeographical region

Species Taxon Far-northern Northern South-eastern Western Elizabeth Overall

Endemic species
Eucidaris galapagensis (Valenciennes) E 54.5)1 264.4 392.9 252.7 280.0 316.8
Octopus oculifer Hoyle M 0 0.02 0.07 0.22 0.18 0.09
Scyllarides astori Holthuis C 0 0 0.06 0.44 0 0.09
Panamic species
Tripneustes depressus (Agassiz) E 0.65)3 45.2 49.11 29.2 2.40)2 37.6
Echinometra vanbrunti (Agassiz) E 0)8 0.66)6 0.79)4 63.24 230.23 30.2
Diadema mexicanum (Agassiz) E 65.41 2.98)3 5.65)3 2.67)3 0.70)4 8.88
Toxopneustes roseus (Agassiz) E 0.07 0.24 0.28 0 0 0.19
Aniculus elegans Stimpson C 0 0.18 0.27 0 0 0.17
Holothuria difficilis Semper H 0 0 0 0.16 0.09 0.03
Holothuria imitans (Ludwig) H 0 0 0.01 0 0 0.01
Trizopagurus magnificus (Bouvier) C 0 0.01 0.01 0 0 0.01
Holothuria impatiens (Forsckal) H 0.02 0 0 0 0 0
Luidia foliata A 0 0 0 0 0.04 0

Indo-Pacific species
Holothuria kefersteini (Selenka) H 1.22 2.95 3.874 0.68)4 2.75 2.96
Holothuria atra (Jaeger) H 0.23)7 3.091 2.345 0.02)5 0.03)5 1.75
Stichopus horrens Selenka H 0.3 0.08 0.72 0.02 0.03 0.43
Holothuria fuscocinerea (Jaeger) H 0.14 0.802 0.38 0 0 0.34
Mithrodia bradleyi Verrill A 0 0.05 0.26 0 0 0.15
Asteropsis carinifera (Lamarck) A 0.02 0.14 0.07 0 0.02 0.06
Panulirus penicillatus (Olivier) C 0.55 0.01 0.02 0 0 0.06

Widespread species
Lytechinus semituberculatus (Agassiz) E 0)2 40.7)1 103.6 369.31 44.3)1 120.6
Stichopus fuscus (Ludwig) H 0.19)4 0.96)2 1.46)1 30.42 32.51 8.33
Hexaplex princeps (Broderip) M 6.45 7.82 9.562 5.91)1 3.32)3 7.96
Pentaceraster cumingi (Gray) A 0)6 4.27 4.833 1.04)2 3.71 3.71
Nidorellia armata (Gray) A 0)5 0.43)4 1.61)2 11.23 11.52 3.60
Centrostephanus coronatus (Verrill) E 0)9 1.25)5 0.46)5 9.355 15.35 3.18
Phataria unifascialis (Gray) A 0 0.08)7 0.21)6 2.216 1.714 0.61
Pharia pyramidata (Gray) A 0 0.02 0.05 0.667 1.746 0.28
Linckia columbiae Gray A 0.02 0 0.03 1.41 0.40 0.26
Percnon gibbesi (H. Milne Edwards) C 2.192 0.10 0 0.01 0 0.18
Pleuroploca princeps Sowerby M 0 0.02 0.05 0.07 0.10 0.05

Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd 1119

G. J. Edgar et al.

(a) 0.2 Far-northern (c) 0.8

Northern
Southeastern 0.6
Abudefduf troschelii
Western Haemulon scudderi
Elizabeth Halichoeres nicholsi
0.1 Anisotremus interruptus
0.4

Johnrandallia nigrirostris
Prionurus laticlavius 0.2
CAP 2

Epinephelus panamensis 0.2 0.4 0.6 0.8

0
0 Zanclus cornutus
Scarus rubroviolaceus
Aulostomus chinensis Heterodontus quoyi
Thalassoma lucasanum Oplegnathus insignis
Kyphosus elegans
Kyphosus analogus Bodianus eclancheri
Semicossyphus darwini
Arothron meleagris
– 0.1 Cirrhitichthys oxycephalus
Melichthys niger
Thalassoma grammaticum
Gymnothorax dovii
Acanthurus nigricans
Heteropriacanthus cruentatus

– 0.2

(b) 0.2 (d)

0.8

0.6

0.1
0.4 Halichoeres dispilus
Ophioblennius steindachneri
Xenocys jessiae
0.2 Labrisomus dendriticus
Holacanthus passer
CAP 3

0 0.2 0.4 0.6 0.8

0
Stegastes arcifrons
Canthigaster punctatissima
Sufflamen verres Mycteroperca olfax
Paralabrax albomaculatus
– 0.1 Lythrypnus gilberti

– 0.2
– 0.2 – 0.1 0 0.1 0.2
CAP 1

Figure 12 Results of CAP analysis maximizing differences in fish data between five bioregions, showing distribution of 0.01 · 0.01
grid cells with CAP axes 1 and 2 (a) and axes 1 and 3 (b). Correlations of species abundance with the three CAP axes are also shown
for species with r > 0.4 (c, d). Species showing high correlations with both CAP axes 2 and 3 are depicted on the graph showing
strongest correlation.

connectivity, and slow recolonization rates when subpopula-
tions have become extinct. A high level of connectivity between
fish populations on different islands is indicated by maximal
fish species richness occurring around the small isolated far-
northern islands of Darwin and Wolf. The only explanation for
this observation within the framework of current island
biogeography theory is that high immigration rates prevail
(Whittaker, 1998). By contrast, macro-invertebrate species
richness was lowest amongst the isolated far-northern islands,
a clear indication that immigration rates are less than for
fishes.
Implications for conservation management
Governments worldwide are increasingly recognizing that
conservation of marine biodiversity requires inter alia a
network of wildlife refuges such as marine protected areas
(MPAs) that are distributed across all biogeographical regions
within their jurisdiction (Australian and New Zealand Envi-
ronment and Conservation Council Task Force on Marine
Protected Areas, 1999; Roff & Taylor, 2000; Roff & Evans,
2002). In most cases, the biogeographical framework for
reserves is defined using geophysical approaches, with regions
classified in terms of abiotic factors such as salinity,
temperature, depth, rock type or sediment particle size that
are considered surrogates for biota (Roff et al., 2003). How-
ever, physical factors rarely act in synchrony, and congruence
between physical and biological regionalizations has rarely
been assessed.
Biological as well as physical regionalizations can vary
greatly, depending on which component of the biota is
analysed. For example, a Tasmanian bioregionalization based
on shallow reef biota exhibited few similarities to one based
on the estuarine fauna (Edgar et al., 1997, 2000). Clearly,
regionalizations used for marine management should reflect
the management target, either directly or by using physical
surrogates shown to covary closely with the subject of
primary interest. In the case of MPA networks, areas are
designated primarily to protect fished species and associated
ecosystems. For this reason, our Galapagos marine bioregio-
nalization was based on inshore reef ecosystems, the habitat
most heavily targeted by local fishers. Sea cucumbers and

1120 Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd

 Regional biogeography of Galapagos shallow-water reef fauna

(a) 0.2 Far-northern (c) 0.8

Northern
Southeastern 0.6
Western Eucidaris galapagensis
0.1 Elizabeth 0.4

Holothuria atra
0.2 Centrostephanus coronatus
CAP 2

0 0.2 0.4 0.6 0.8

0
Echinometra vanbrunti
Phataria unifascialis

Percnon gibbesi
– 0.1 Panulirus penicillatus

Diadema mexicanum

– 0.2

(b) 0.1
(d) 0.8
Lytechinus semituberculatus
0.6

Tripneustes depressus 0.4

CAP 3

0.2

0 0.4 0.2 0
Holothuria difficilis

Pharia pyramidata
Nidorellia armata
Holothuria fuscocinerea
Stichopus fuscus
Pentaceraster cumingi

– 0.1
– 0.2 – 0.1 0 0.1 0.2
CAP 1

Figure 13 Results of CAP analysis maximizing differences in macro-invertebrate data between five bioregions, showing distribution of
0.01  · 0.01  grid cells with CAP axes 1 and 2 (a) and axes 1 and 3 (b). Correlations of species abundance with the three CAP axes are also
shown for species with r > 0.4 (c, d). Species showing high correlations with both CAP axes 2 and 3 are depicted on the graph showing
strongest correlation.

Table 3 Classification of fish data pertain-

Region Far-northern Northern South-eastern Western Elizabeth Total Correct (%)
ing to 0.01 · 0.01 grid cells to five
biogeographical regions using the leave-
Far-northern 6 0 0 0 0 6 100
one-out procedure of canonical analysis of
Northern 1 20 3 1 0 25 80
principal coordinates
South-eastern 0 14 108 3 2 127 85
Western 0 0 4 31 6 41 76
Elizabeth 0 0 3 5 13 21 62

Table 4 Classification of invertebrate data

Region Far-northern Northern South-eastern Western Elizabeth Total Correct (%)
pertaining to 0.01 · 0.01 grid cells to five
biogeographical regions using the leave-
Far-northern 5 1 0 0 0 6 0.83
one-out procedure of canonical analysis of
Northern 0 15 8 0 0 23 0.65
principal coordinates
South-eastern 1 28 77 4 2 112 0.69
Western 0 0 1 26 9 36 0.72
Elizabeth 0 0 0 3 11 14 0.79

lobsters captured by divers provided > 80% of the total Conservation of marine biodiversity in Galapagos would be
annual income of fishers over the past 5 years (Murillo, assisted by a change in the GMR Management Plan from
2002). recognition of the Harris biogeographical regions to the five

Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd 1121

G. J. Edgar et al.

Table 5 Area (km2) included in different management zones (Valle, 1995), Galapagos penguin (Boersma, 1998) and
for the five biogeographical regions defined in this study Galapagos fur seal (Trillmich & Limberger, 1985).
Management zone, km2 (%)
Biogeographical
ACKNOWLEDGMENTS
region Conservation Tourism Fishing Total
We would like to thank the various divers who assisted us
Far-northern 0.78 (4.6) 3.78 (22.2) 12.46 (73.2) 17.02
collecting data, particularly Lauren Garske, Noemi D’Ozou-
Northern 6.23 (5.5) 18.25 (16.0) 89.46 (78.5) 113.94
ville, Linda Kerrison, Fernando Rivera, Vicente Almardariz,
South-eastern 68.88 (7.7) 98.92 (11.0) 731.49 (81.3) 899.29
Veronica Toral-Granda, Scoresby Shepherd, Giancarlo Toti,
Western 36.61 (11.2) 26.37 (8.1) 262.93 (80.7) 325.91
Elizabeth 7.68 (4.8) 16.80 (10.6) 134.64 (84.6) 159.12 Julio Delgado, Angel Chiriboga, Diego Ruiz and Vanessa
Francesco. Support for the monitoring program provided by
Total 120.17 164.12 1230.98 1515.27
Galapagos National Park Service, and funding by USAID, the
Charles Darwin Foundation Inc., Galapagos Conservation
Trust, the British Embassy, Beneficia Foundation, Pew Char-
biogeographical regions outlined here. During consensual itable Trust and Rockefeller Foundation is gratefully acknow-
discussions in 2000, stakeholders of the GMR agreed on ledged. We also thank Rodrigo Bustamante and Robert
preliminary ‘no-take’ conservation zones to protect coastal Bensted-Smith for organizing funding proposals, and Marti
ecosystems within each of the five Harris zones; however, Anderson for statistical advice on using the CAP program.
disproportionately low levels of protection have consequently
been afforded to the small distinctive ecosystems in the far-
REFERENCES
northern and Elizabeth bioregions.
The far-northern region has very high conservation signi- Anderson, M.J. (2003) CAP: a FORTRAN computer program
ficance due to its distinctive biota with anomalously high for canonical analysis of principal coordinates. Department of
species richness of fishes and corals. The area reserved for Statistics, University of Auckland, Auckland, New Zealand.
conservation purposes within this bioregion is very small Anderson, M.J. & Robinson, J. (2003) Generalised dis-
(0.78 km2, Table 5), in part because the bioregion itself is so criminant analysis based on distances. Australian and New
small, comprising two islands only. Zealand Journal of Statistics, 45, 301–318.
The Elizabeth bioregion includes the smallest percentage of Anderson, M.J. & Willis, T.J. (2003) Canonical analysis of
coast zoned for conservation amongst the five Galapagos principal coordinates: a useful method of constrained or-
biogeographical regions described here (Table 5), with only a dination for ecology. Ecology, 84, 511–524.
single small area around Islas Marielas fully protected. A Australian and New Zealand Environment and Conservation
notable feature of this bioregion is a disproportionately high Council Task Force on Marine Protected Areas (1999)
number of endemic Galapagos species, making it the core area Strategic plan of action for the National Representative System
for much of the endemic Galapagos inshore marine fauna and of Marine Protected Areas: a guide for action by Australian
possibly an important refuge for local Galapagos species during Governments. Environment Australia, Canberra.
periods of adverse environmental conditions such as El Niño. Banks, S.J. (2002) Ambiente Fı́sico. Reserva Marina de Gal-
Thus, despite its low overall species richness, the Elizabeth ápagos, Lı́nea Base de la Biodiversidad (ed. by E. Danulat and
bioregion should be considered an area with exceptional G.J. Edgar), pp. 18–33. Charles Darwin Foundation and
conservation significance. Galápagos National Park Service, Galápagos, Ecuador.
The inverse relationship evident at regional scales in Bensted-Smith, R. (2002) A biodiversity vision for the Galapagos
Galapagos between the richness of the total fish fauna and Islands. Charles Darwin Foundation and World Wildlife
the richness of endemic fish species has broad management Fund, Puerto Ayora, Galapagos, Ecuador.
implications. In the terrestrial environment, several authors Boersma, P.D. (1998) Population trends of the Galapagos
have suggested that conservation areas should be prioritized penguin: impacts of El Nino and La Nina. Condor, 100, 245–
spatially in terms of ‘hotspots’ of biodiversity (Meyers et al., 253.
2000). Roberts et al. (2002) extended this concept to the Bustamante, R., Collins, K.J. & Bensted-Smith, R. (2000)
marine realm, arguing that centres of marine endemism in the Biodiversity conservation in the Galápagos Marine Reserve.
Pacific are congruent with hotspots of species richness (but for Bulletin de l’Institut Royal des Sciences Naturelles de Belgigue,
alternate view see Hughes et al., 2002). A hotspot strategy for Supplement 70, 31–38.
Galapagos, in the absence of gap analysis, would indicate a low Bustamante, R.H., Vinueza, L.R., Smith, F., Banks, S., Calv-
conservation rating for western Galapagos and Elizabeth opiña, M., Francisco, V., Chiriboga, A. & Harris, J. (2002)
ecosystems, whereas on a global scale these areas possess Comunidades submareales rocosas. I: Organismos sésiles y
highest conservation significance. The western upwelling mesoinvertebrados móviles. Reserva Marina de Galápagos,
region provides core habitat for virtually all endemic Galap- Lı́nea Base de la Biodiversidad (eds E. Danulat and G.J.
agos marine taxa, including many charismatic species not Edgar), pp. 33–62. Charles Darwin Foundation and
included in the present study, such as the flightless cormorant Galápagos National Park Service, Galápagos, Ecuador.

1122 Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd

 Regional biogeography of Galapagos shallow-water reef fauna
Cairns, S.D. (1991) A revision of the ahermatypic Scleractinia
of the Galápagos and Cocos Islands. Smithsonian Contri-
butions in Zoology, 504, 1–32.
Carr, M.R. (1996) PRIMER User Manual. Plymouth Routines in
Multivariate Ecological Research. Plymouth Marine Labora-
tory, Plymouth, UK.
Clarke, K.R. (1993) Non-parametric multivariate analyses of
changes in community structure. Australian Journal of
Ecology,18, 117–143.
Edgar, G.J., Moverley, J., Barrett, N.S., Peters, D. & Reed, C.
(1997) The conservation-related benefits of a systematic
marine biological sampling program: the Tasmanian reef
bioregionalisation as a case study. Biological Conservation,
79, 227–240.
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 Con-
servation, 92, 383–397.
Faith, D.P., Minchin, P.R. & Belbin, L. (1987) Compositional
dissimilarity as a robust measure of ecological distance.
Vegetatio, 69, 57–68.
Glynn, P.W. (1994) State of coral reefs in the Galapagos Is-
lands: natural vs anthropogenic impacts. Marine Pollution
Bulletin, 29, 131–140.
Glynn, P.W. & Ault, J.S. (2000) A biogeographic analysis and
review of the far eastern Pacific coral reef region. Coral Reefs,
19, 1–23.
Grove, J.S. (1985) Influence of the 1982–1983 El Niño event
upon the ichthyofauna of the Galápagos archipelago. El Niño
in the Galápagos Islands: the 1982–1983 Event (ed. by G.
Robinson and E.M. Del Pino), pp. 191–198. Charles Darwin
Foundation for the Galápagos Islands, Quito, Ecuador.
Harris, M.P. (1969) Breeding season of sea-birds in the Gal-
apagos Islands. Journal of Zoology, 159, 145–165.
Hickman, C.P. (1998) A field guide to sea stars and other
echinoderms of Galápagos. Sugar Spring Press, Lexington,
MA, USA.
Houvenaghel, G.T. (1978) Oceanographic conditions in the
Galápagos Archipelago and their relationships with life on
the islands. Upwelling Ecosystems (eds R. Boje and M.
Tomczak), pp. 181–200. Springer-Verlag, Berlin.
Hughes, T.P., Bellwood, D.R. & Connolly, S.R. (2002) Biodi-
versity hotspots, centres of endemicity, and the conservation
of coral reefs. Ecology Letters, 5, 775–784.
Humann, P. (1988) Galápagos – Un phenómeno terrestre y
marino. Libri-Mundi, Quito, Ecuador.
Jennings, S., Brierley, A.S. & Walker, J.W. (1994) The inshore
fish assemblages of the Galapagos Archipelago. Biological
Conservation, 70, 49–57.
Lessios, H.A., Kessing, B.D., Robertson, D.R. & Paulay, G.
(1999) Phylogeography of the pantropical sea urchin Eu-
cidaris in relation to land barriers and ocean currents.
Evolution, 53, 806–817.
Meekan, M.G., Ackerman, J.L. & Wellington, G.M. (2001)
Demography and age structures of coral reef damselfishes in
Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd
the tropical eastern Pacific Ocean. Marine Ecology Progress
Series, 212, 223–232.
Myers, N., Mittermeier, R.A., Mittermeier, C.G., da Fonseca,
G.A. & Kent, J. (2000) Biodiversity hotspots for conserva-
tion priorities. Nature, 403, 853–858.
Murillo, J.C. (2002) Indicadores socio-económicos de la pesca.
Reserva Marina de Galápagos, Lı́nea Base de la Biodiversidad
(ed. by E. Danulatand G.J. Edgar), pp. 440–448. Charles
Darwin Foundation and Galápagos National Park Service,
Galápagos, Ecuador.
Pak, H. & Zanfield, J.R.V. (1974) The Cromwell Current on
the east side of the Galápagos. Journal of Geophysical Re-
search, 78, 7845–7859.
Roberts, C.M., McClean, C.J., Veron, J.E.N., Hawkins, J.P.,
Allen, G.R., McAllister, D.E., Mittermeier, C.G., Schueler,
F.W., Spalding, M., Wells, F., Vynne, C. & Werner, T.B.
(2002) Marine biodiversity hotspots and conservation
priorities for tropical reefs. Science, 295, 1280–1284.
Robinson, G. (1985) Influence of the 1982–83 El Niño on
Galápagos marine life. El Niño in the Galápagos Islands: the
1982–1983 event (ed. by G. Robinson and E.M. Del Pino),
pp. 153–190. Charles Darwin Foundation for the Galápagos
Islands, Quito, Ecuador.
Roff, J.C. & Evans, S. (2002) Frameworks for marine con-
servation – non-hierarchical approaches and distinctive
habitats. Aquatic Conservation: Marine and Freshwater Eco-
systems, 12, 635–648.
Roff, J.C. & Taylor, M.E. (2000) National frameworks for marine
conservation – a hierarchical geophysical approach. Aquatic
Conservation: Marine and Freshwater Ecosystems, 10, 209–223.
Roff, J.C., Taylor, M.E. & Laughren, J. (2003) Geophysical
approaches to the classification, delineation and monitoring
of marine habitats and their communities. Aquatic Con-
servation: Marine and Freshwater Ecosystems, 13, 77–90.
Trillmich, F. & Limberger, D. (1985) Drastic effects of El Niño
on Galápagos pinnipeds. Oecologia, 67, 19–22.
Valle, C. (1995) Effective populations size and demography of
the rare flightless Galápagos cormorant. Ecological Applica-
tions, 5, 601–617.
Victor, B.C., Wellington, G.M., Robertson, D.R. & Ruttenberg,
B.I. (2001) The effect of the El Niño-Southern Oscillation
event on the distribution of reef-associated labrid fishes in the
eastern Pacific Ocean. Bulletin of Marine Science, 69, 279–288.
Wellington, G.M. (1975) The Galápagos coastal marine envi-
ronment. Resource report, Department of National Parks
and Wildlife, Quito, Ecuador.
Wellington, G.M. & Victor, B.C. (1992) Regional differences in
duration of the planktonic larval stage of reef fishes in the
eastern Pacific Ocean. Marine Biology, 113, 491–498.
Wellington, G.M., Strong, A.E. & Merlen, G. (2001) Sea sur-
face temperature variation in the Galapagos Archipelago: a
comparison between AVHRR night-time satellite data and
in situ instrumentation (1982–1998). Bulletin of Marine
Science, 69, 27–42.
Whittaker, R.J. (1998) Island biogeography: ecology, evolution,
and conservation. Oxford University Press, Oxford.
1123
G. J. Edgar et al.

BIOSKETCHES

Graham Edgar recently returned to Tasmania after 2 years as Head of Marine Science at the Charles Darwin Research Station,
Galapagos. He has investigated a variety of ecological interactions in the marine environment, primarily those involving human
impacts.

Stuart Banks is mainly interests in physical oceanography, particularly the interpretation of oceanographical patterns at large
spatial scales using satellite imagery.

José Miguel Fariña currently holds a faculty position at the Center for Advanced Studies in Ecology and Biodiversity, Santiago,
after returning from postdoctoral studies at Brown University. His research has focussed on ecotrophic transfer between land and sea
and impacts of human activity on fishes and benthos.

Mónica Calvopiña is primarily interested in community ecology and the social implications of marine research, particularly as they
affect inhabitants of her Galapagos birthplace.

Camilo Martı́nez recently completed a masters degree in marine resource management in the Canary Islands. He has worked
extensively on the biology of fishery species, most notably the Galapagos spiny lobster.

1124 Journal of Biogeography 31, 1107–1124, ª 2004 Blackwell Publishing Ltd