Aquatic Botany 121 (2015) 26–32

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Aquatic Botany
journal homepage: www.elsevier.com/locate/aquabot

Effects of bottom topography and anthropogenic pressure on

northern Adriatic Cystoseira spp. (Phaeophyceae, Fucales)
Massimo Devescovi ∗
Rud̄er Bošković Institute, Center for Marine Research, G. Paliaga 5, 52210 Rovinj, Croatia

a r t i c l e i n f o a b s t r a c t

Article history: The effect of bottom topography on Cystoseira spp. was estimated along the west Istrian Coast (northern
Received 6 March 2014 Adriatic Sea, Croatia) between 6 and 10 m depth in off-port control areas and in ports of coastal cities.
Received in revised form 23 October 2014 Two geomorphologic bottom types were considered: a stepped rocky bottom (T1, slope ≈ 10◦ ) and a flat
Accepted 27 October 2014
rocky-sandy bottom (T2, slope ≈ 4◦ ). In control areas, Cystoseira spp. formed mixed stands composed of
Available online 13 November 2014
six species. The bottom topography strongly influenced C. corniculata (dominant on T1) and C. foeniculacea
(dominant on T2). The response of C. barbata and C. spinosa to bottom topography varied among control
Keywords:
areas, whereas C. compressa and C. humilis did not show topographic preferences. In ports, stands were
Cystoseira
Topography
mainly composed of C. compressa and C. humilis. The species C. barbata, C. corniculata, C. foeniculacea and
Anthropogenic pressure C. spinosa were strikingly regressed, leading to a decrease in Cystoseira spp. total cover. However, they
Port activities tended to be more frequent on T1 than on T2, which highlighted the role of bottom topography for their
Rocky bottom survival in adverse conditions. The capability of C. compressa and C. humilis to withstand moderate levels
Adriatic Sea of anthropogenic pressure that is characteristic of west Istrian ports may well be related to their rapid
growth and high recruitment rate.
© 2014 Elsevier B.V. All rights reserved.

1. Introduction
Species of the genus Cystoseira (Phaeophyceae, Fucales) are large
and long-lasting canopy-forming brown macroalgae. A total of 41
taxa (at the species and infraspecific level) are currently reported
from the Mediterranean Sea (Taşkin et al., 2012). They are usu-
ally firmly attached to the rocky substrate by a basal disc and are
widespread from the shallow coastal areas to over 100 m depth,
forming mono-specific or mixed stands that vary in species compo-
sition among Mediterranean biogeographic sectors and with depth
(Ercegović, 1952; Giaccone and Bruni, 1973). The complex struc-
ture of their thalli provides suitable habitats for a large number of
algal and animal species, thus sustaining high biodiversity levels
of coastal ecosystems (Falace and Bressan, 2006; Ballesteros et al.,
1998, 2009).
Urban and industrial pollution can cause local extinction of Cys-
toseira spp. (Mangialajo et al., 2008; Sales et al., 2011). At a larger
spatial scale, eutrophication, related to increased anthropogenic
alteration of biogeochemical cycles and global warming (Boyd and
Hutchins, 2012), leads to light limitation that adversely affects the
abundance of benthic primary producers (Kavanaugh et al., 2009)
∗ Tel.: +385 52 804 73; fax: +385 52 804 780.
E-mail address: massimo.devescovi@cim.irb.hr
and hence Cystoseira spp. populations (Munda, 1993; Cormaci and
Furnari, 1999). Given their sensitivity to anthropogenic pressure,
Cystoseira spp. have been used as biological indicators of the eco-
logical quality of coastal waters (Panayotidis et al., 2004; Ballesteros
et al., 2007; Juanes et al., 2008) under the requirements of the
European Water Framework Directive (WFD; EC, 2000).
However, other human-induced and natural processes affect
Cystoseira spp. populations. Sea urchin grazing has been shown
to markedly alter the underwater landscape, converting stands of
large erect algae into barrens; one of the factors leading to increased
sea urchin densities could be the overfishing of their predators
(Sala et al., 1998; Shears and Babcock, 2002). Another impact is the
worldwide documented extirpation of brown algae during strong
storms (Gunnill, 1986; Underwood, 1998; Pardi et al., 2000). Dele-
terious effects of exceptional storms on Cystoseira spp. stands were
observed up to a depth of approximately 20 m (Navarro et al., 2011).
It could be suggested that the reshaping of Cystoseira spp.
assemblage composition has emerged from interactions among
external perturbing factors and patterns of myriad biological and
hydrodynamic processes inherent to each habitat. Most of these
processes are related to habitat geomorphology, which acts as an
essential environmental factor contributing to the structure and
function of benthic communities (Tokeshy and Arakaki, 2012). Geo-
morphology also influences predation rates (Fairweather, 1988;
Johnson et al., 1998) and larval settlement (Idjadi and Edmunds,

http://dx.doi.org/10.1016/j.aquabot.2014.10.009
0304-3770/© 2014 Elsevier B.V. All rights reserved.
 M. Devescovi / Aquatic Botany 121 (2015) 26–32
Fig. 1. A map of the study area. P1, P2 and P3 are randomly selected ports. C1, C2
and C3 are randomly selected control areas.
2006) and provides refuge against physical and biological distur-
bances (Menge et al., 1985; Bergeron and Bourget, 1986). Previous
studies have demonstrated the importance of some bottom topo-
graphic variables such as the bottom slope (Díez et al., 2003), the
percent of rocky bottom extent (Sales and Ballesteros, 2009) and the
architecture of rocky formations (Eriksson and Bergström, 2005;
Simakova, 2009; Wallin et al., 2011) on the demography of some
Cystoseira spp. and other members of the order Fucales.
The aim of this study was to compare the abundance of indi-
vidual Cystoseira spp. between off port control areas and ports of
coastal cities, separating the effects of anthropogenic pressure and
bottom topography by a factorial model strategy. The consequences
of anthropogenic pressure on Cystoseira spp. distribution have
typically been assessed at depths between 0 and several metres.
However, in ports, the natural topography at these depths is usually
radically modified (e.g., by dredging and construction of artificial
structures). Hence, to allow a comparison between impacted and
control areas for habitats of similar topography, sampling was per-
formed from 6 to 10 m depth. The study area was the west coast
of the Istrian Peninsula located in the northern Adriatic Sea, which
represents the northernmost biogeographic sector of the Mediter-
ranean Sea (Bianchi and Morri, 2000). In contrast to other northern
Adriatic regions, the sea bottom along the Istrian Coast is mainly
rocky and thus particularly adapted to harbour Cystoseira spp.
populations.
2. Methods
2.1. Study area
The west Istrian Coast (90 km by a straight line transect from
Cape Savudrija to Cape Kamenjak, Fig. 1) represents 17% of the
total Croatian coast (by a straight line transect, 526 km); it is a
very important tourist destination sustaining almost 35% of the
total Croatian tourism sector. Three ports of coastal cities, encom-
passing approximately 2.5 km of the coast each (by a straight
line transect), were randomly selected (Umag = P1, Poreč = P2 and
27
Fig. 2. The topographic variables assessed for rocky bottom profiles. The rocky
substrate is shown in black; empty segments (s1 , s2 , . . . sn ) are lengths of sandy
patches. The lengths of the approximately vertical rocky substrate are indicated by
h1 , h2 , . . . hn . The profile slope = tg˛; the sandy bottom fraction = S%; and the verti-
cal relief = H%. The depth range (a) was fixed (4 m); the distance range (b) varied
according to the slope. The length of the hypotenuse (c) was calculated by simple
trigonometry.
Rovinj = P3, Fig. 1). Each port includes marinas (200–800 moorings),
fishing boat harbours and harbours for the boats of local inhabi-
tants. Yachts and cruise liners anchor in ports during the tourism
season. The number of inhabitants in the cities is between 7000 and
11,000; however, this number increases several times during the
tourism season. Control areas C1, C2 and C3 (Fig. 1) were randomly
selected from natural bays with a coastal morphology and wave
exposure patterns comparable to those of ports. They are also likely
subject to slight anthropogenic pressure originating from camping
areas for tourists (10–30% of the coast line) and agriculture (vine-
yards and olive groves, 10–40% of the coast line). Mediterranean
forest covers the remaining coastline of these bays.
2.2. Bottom topography
Geometrical features of the bottom profile were assessed
according to Devescovi et al. (2005) along a transect perpendicular
to the coast from 6 to 10 m depth. From the obtained data, values
of selected topographic variables (slope, sandy bottom fraction and
the percent of vertical rocky relief) were calculated (Fig. 2). Two
types of bottom topography were considered for Cystoseira spp.
sampling: a stepped rocky bottom (T1; slope = 8–12◦ , sandy bottom
fraction = 7–20% and vertical relief = 12–22%) and a flat rocky-sandy
bottom (T2; slope = 3–5◦ , sandy bottom fraction = 47–63% and ver-
tical relief = 2–3%).
2.3. Cystoseira spp. sampling
Preliminary observations showed that in sampled habitats, Cys-
toseira spp. formed mixed stands composed of six species: (1) C.
barbata (Stackhouse) C. Agardh, (2) C. compressa (Esper) Gerloff and
Nizamuddin, (3) C. corniculata (Turner) Zanardini, (4) C. foeniculacea
(Linnaeus) Greville, (5) C. humilis Schousboe ex Kützing and (6) C.
spinosa Sauvageau.
The sampling was performed at a depth of 6–10 m in spring
(April 2009). Cystoseira spp. deciduous branches were well
developed during this period, which permitted underwater iden-
tification of thalli (SCUBA diving). Cystoseira spp. abundance was
assessed underwater as areal cover (i.e., the area of the substrate
covered by an orthogonal projection of thalli). The cover was
estimated using a 50 cm × 50 cm quadrat subdivided into 9 sub-
squares. This subdivision was chosen because of the large size of
thalli. Within each sub-square, the cover was estimated as a pro-
portion: 1/4, 2/4, 3/4 and 4/4. If less than 1/4 of the square was
28 M. Devescovi / Aquatic Botany 121 (2015) 26–32

Table 1
A four-factor ANOVA for Cystoseira spp. total cover.

Source DF MS F P F-denominator

I 1 246.257 47.652 0.002 A(I)

T 1 0.909 0.953 0.384 T × A(I)
I×T 1 2.033 2.131 0.218 T × A(I)
A(I) 4 5.168 13.168 <0.001 Site(T × A(I))
T × A(I) 4 0.954 2.431 0.105 Site(T × A(I))
Site(T × A(I)) 12 0.392 0.664 0.781 Residual
Residual 96 0.591

Transformation: square root. Levene’s test: P = 0.474

Tukey’s HSD tests for A(I)

Control areas: C1 < C3 = C2; Ports: P1 = P3 = P2

Factors are: I = impact, fixed, 2 levels: control areas and ports; T = topography, fixed, 2 levels: the stepped rocky bottom (T1) and the flat rocky-sandy bottom (T2); A(I) = area
nested in I, random, 3 levels: 3 control areas (C1, C2 and C3) and 3 ports (P1, P2 and P3); Site(T × A(I)) = site nested in T × A(I), random, 2 levels. Number of replicates: n = 5;
N = 120.

filled, an arbitrary value of 1/10 was assigned. The proportions of
all squares were summed, and the resulting value was multiplied
by 100/9 to obtain the percent cover of the quadrat. To allow sta-
tistical comparison of cover among species, Cystoseira spp. were
sampled independently (i.e., the cover of only one species was
assessed within a given quadrat; Underwood, 1997). At each site, 5
randomly placed quadrats were sampled for each species. The total
cover of all Cystoseira spp. present in a quadrat was also assessed at
each site for 5 randomly placed quadrats. Within each control area
and within each port, two sites were randomly chosen for each bot-
tom type so that the total number of replicates for each species and
total cover was 120.
In ports, the cover of C. compressa and C. humilis was particularly
high. This could be due to rapid growth and/or a high recruitment
rate of these two species. To investigate these patterns, the size of
thalli and the length of the perennial axes of C. compressa and C.
humilis collected in ports were compared with those of all species
found the in control areas. The biomass of thalli was used as a
measure of size; the length of the axes was considered as an approx-
imation of age (Ercegović, 1952; Ballesteros et al., 2009). At each
site, 5 thalli for each species considered were randomly collected.
In the laboratory, the wet weight of each thallus was determined
after blotting the adhering sea water by filter paper. The length of
the main axis was assessed for C. barbata and C. spinosa thalli. Thalli
of C. compressa, C. foeniculacea and C. humilis are caespitose (more
main axes per thallus), so the length of the longest axis was mea-
sured. For C. corniculata, the total weight of all perennial parts of
each thallus was assessed. In this species, main axis length assess-
ment can hardly be accomplished; the main axis grows horizontally
to the substrate and produces many secondary axes forming an
intricate and compact aggregate.
Model 3: C. barbata, C. corniculata, C. foeniculacea and C. spinosa
abundances were compared among control areas. The factors were
Species (fixed, 4 levels), Topography (as in model 1), Area (random,
3 levels) and Site (random and nested in the interaction Topogra-
phy × Area, 2 levels). The dependent variable was cover.
Data for the wet weight of thalli and the length of axes
were graphically explored using box-and-whisker plots. Differ-
ences among sample medians were assessed using a multi-sample
median test followed by Tukey-type multiple comparison tests for
medians (Zar, 1999).
3. Results
3.1. Cystoseira spp. total cover
Total cover was significantly lower in ports than in control areas
(Table 1 and Fig. 3). Factor Area (Impact) was significant showing
that there were differences in total cover among randomly selected
areas within Impact levels. In control area C1, total cover was sig-
nificantly lower than in control areas C2 and C3. The total cover
did not differ among ports. Differences between topographic types
were not detected both within and across areas (Table 1).

2.4. Data analysis

Depending on the structure of the data, analyses of variance

(ANOVA) were performed according to three linear models using a
software package SYSTAT 12 (SYSTAT Software, Inc.).

Model 1: The model was composed of four factors: Impact (fixed,

2 levels: port and control), Topography (fixed, 2 levels: T1 and
T2), Area (random and nested in Impact, 3 levels: 3 ports and 3
control areas were randomly chosen along the west Istrian Coast)
and Site (random and nested in the interaction Topography × Area
(Impact), 2 levels: 2 randomly chosen sites within each combina-
tion of Topography and Area levels). The dependent variable was
the total Cystoseira spp. cover.
Model 2: The above four factors were included in the model with Fig. 3. The total cover of all Cystoseira spp. assessed in three control areas (C1, C2
and C3), three ports (P1, P2 and P3), and two different bottom topographic types.
the addition of the factor Species (fixed, 2 levels: C. compressa and The empty bars indicate a stepped rocky bottom (T1); the filled bars indicate a flat
C. humilis). The dependent variable was cover. rocky-sandy bottom (T2). The data are means ± SD, n = 10.
 M. Devescovi / Aquatic Botany 121 (2015) 26–32
Fig. 4. The cover of individual Cystoseira spp. in three control areas (a) and three
ports (b). Empty and filled bars indicate a stepped rocky bottom and flat rocky-
sandy bottom, respectively. Abbreviations: barb = C. barbata; comp = C. compressa;
corn = C. corniculata; foen = C. foeniculacea; humi = C. humilis; spin = C. spinosa. The
data are means ± SD, n = 10. The other details are the same as in Fig. 3.
3.2. Distribution of Cystoseira compressa and Cystoseira humilis
An ANOVA comparing the cover of C. compressa and C. humilis
detected the significance of the interaction Species × Impact. This
indicated that the main factors Species and Impact were not inde-
pendent; consequently, individual tests for these two factors were
unreliable and their effects need to be interpreted in the framework
of the interaction (Underwood, 1997). The results of unplanned
comparisons (Tukey’s HSD tests, Table 2) for this two-factor inter-
action can be summarised in four points. (1) In control areas, C.
humilis was more abundant than C. compressa. (2) In ports, C. com-
pressa was more abundant than C. humilis. (3) C. compressa was
more abundant in ports than in control areas. (4) The abundance
of C. humilis in control areas was similar to that in ports. The cover
values of both species for each area are reported in Fig. 4.
Factor Area (Impact) was significant (Table 2), indicating dif-
ferences among areas within Impact levels (means obtained by
averaging across species and bottom types). Significant effects of
bottom topography on the cover of C. compressa and C. humilis were
not detected.
3.3. Distribution of the other Cystoseira spp.
In ports, C. corniculata and C. spinosa were not found on the
bottom type T2, whereas low amounts of these two species were
noticed on T1. Few C. barbata and C. foeniculacea were found on both
bottom types (Fig. 4). However, their cover was higher on T1 than on
T2 (one-tailed two-sample t-test for separate variances with n = 30
for C. barbata: t = 2.763, DF = 37.43, P = 0.004 and for C. foeniculacea:
t = 3.438, DF = 48.25, P = 0.001).
ANOVA comparing the cover of C. barbata, C. corniculata, C.
foeniculacea and C. spinosa among control areas detected the
29
Fig. 5. Box-plots for thallus wet weight (a) and axis length (b) of Cysoseira spp.
Samples compP and humiP were collected in ports; the other samples were collected
in control areas. The axis length for C. corniculata was not assessed. Replicates per
sample: n = 60. The other details are the same as in Fig. 4.
significance of the interaction Species × Topography × Area
(Table 3). Consequently, tests for higher order terms were not
considered. The three-factor interaction was examined in detail
by a series of Tukey’s HSD tests (Table 4). The obtained results
can be summarised in four points. (1) Comparisons of species
cover within topography levels revealed that in all control areas, C.
corniculata and C. foeniculacea were the more abundant species on
T1 and T2, respectively. (2) The ranking of C. barbata and C. spinosa
in mixed stands and their relation to bottom topography differed
from area to area. (3) Comparisons of species cover between
topography levels revealed that in all control areas, C. corniculata
was more abundant on T1 than on T2 and that C. foeniculacea was
more abundant on T2 than on T1. (4) The cover of the four species
varied significantly from area to area on both topographic types
except in the case of C. barbata on T1. The cover values of all four
species for each area are reported in Fig. 4.
3.4. Weight of thalli and axis length
The distributions of thallus wet weight for all six species col-
lected in the control areas were graphically compared with those
for C. compressa and C. humilis collected in ports using box-and-
whisker plots (Fig. 5a). The maximum values of thallus weight
increased in the following order: C. foeniculacea (65 g), C. humilis
in ports (75 g), C. compressa (77 g), C. compressa in ports (78 g),
C. humilis (94 g), C. spinosa (152 g), C. barbata (213 g) and C. cor-
niculata (623 g). The median for C. corniculata (145.50 g) largely
exceeded the medians of all other samples. For the other species,
medians ranged from 23.80 (C. humilis) to 42.30 g (C. spinosa). A
multi-sample median test (C. corniculata excluded) did not detect
significant differences (2 = 10.553, DF = 6, P = 0.104, n = 60, N = 420).
30 M. Devescovi / Aquatic Botany 121 (2015) 26–32

Table 2
A five-factor ANOVA comparing cover of Cystoseira compressa and Cystoseira humilis in ports and control areas.

Source DF MS F P F-denominator

Species = S 1 3.597 7.346 0.054 S × A(I)

Impact = I 1 18.409 7.008 0.057 A(I)
Topography = T 1 2.913 6.387 0.065 T × A(I)
S×I 1 48.956 99.986 0.001 S × A(I)
S×T 1 2.124 4.293 0.107 S × T × A(I)
I×T 1 2.354 5.161 0.086 T × A(I)
S×I×T 1 0.594 1.201 0.335 S × T × A(I)
Area(I) = A(I) 4 2.627 6.852 0.004 Site(T × A(I))
S × A(I) 4 0.490 1.855 0.183 S × Site(T × A(I))
T × A(I) 4 0.456 1.190 0.364 Site(T × A(I))
S × T × A(I) 4 0.495 1.874 0.180 S × Site(T × A(I))
Site(T × A(I)) 12 0.383 0.688 0.762 Residual
S × Site(T × A(I)) 12 0.264 0.474 0.928 Residual
Residual 192 0.557

Transformation: square root. Levene’s test: P = 0.118

Tukey’s HSD tests for S × I

C: comp < humi comp: C < P
P: humi < comp humi: P = C

Factor S is fixed, 2 levels: C. compressa = comp and C. humilis = humi. Factors I, T, A(I) and Site(T × A(I)) are as in Table 1. C and P are levels of factor I (control and port). Number
of replicates: n = 5; N = 240. Because the S × I interaction is significant, tests for S and I are unreliable; their F and P values are shown in italics.

Table 3
A four-factor ANOVA comparing Cystoseira barbata, Cystoseira corniculata, Cystoseira foeniculacea and Cystoseira spinosa cover among control areas.

Source DF MS F P F-denominator

Species = S 3 85.064 15.395 0.003 S×A

Topography = T 1 2.518 2.065 0.287 T×A
Area = A 2 15.138 41.273 <0.001 Site(T × A)
S×T 3 56.796 14.412 0.004 S×T×A
S×A 6 5.526 12.714 <0.001 S × Site(T × A)
T×A 2 1.219 3.325 0.107 Site(T × A)
S×T×A 6 3.941 9.068 <0.001 S × Site(T × A)
Site(T × A) 6 0.367 0.316 0.316 Residual
S × Site(T × A) 18 0.435 0.375 0.375 Residual
Residual 192 1.159

Transformation: square root. Levene’s test: P = 0.298

The fixed factor S has 4 levels: C. barbata = barb, C. corniculata = corn, C. foeniculacea = foen and C. spinosa = spin. Factor T is as in Table 1. Factor A is random with 3 levels: C1,
C2 and C3. Factor Site(T × A) with 2 levels is random and nested in the interaction T × A. Number of replicates: n = 5; N = 480. Because the S × T × A interaction is significant,
individual tests for all terms included in the interaction are unreliable; their F and P values are shown in italics.

The distributions of the axis length for all species (except C. restricted to depths less than 3 m. The other historically recorded
corniculata) collected in control areas and for C. compressa and C. six species have been found by me in noticeable quantities off ports
humilis collected in ports are graphically presented in Fig. 5b. For C. between 6 and 10 m depth in April 2009, indicating a good preser-
corniculata, the weight of thalli was linearly related to the weight of vation status of Cystoseira spp. populations in the study area.
perennial parts. Perennial parts amounted to approximately 50% of
the weight of thalli (Fig. 6). The maximum values of axis length
increased in the following order: C. compressa = C. compressa in
ports = C. humilis in ports (3.5 cm), C. humilis (3.7 cm), C. foeniculacea
(17.5 cm) and C. barbata = C. spinosa (23.0 cm). The medians for C.
barbata (10.10 cm), C. foeniculacea (6.00 cm) and C. spinosa (7.45 cm)
largely exceeded those of C. compressa = C. compressa in ports = C.
humilis in ports (1.20 cm) and C. humilis (1.35 cm). The medians of
C. humilis in control areas and in ports did not differ (two sam-
ple median test: 2 = 1.212, DF = 1, P = 0.271, n = 60, N = 120). Hence,
they also did not differ from those of C. compressa in both control
areas and ports.

4. Discussion

At the end of the 19th century, seven Cystoseira spp. were

reported for the region of Rovinj in Paul Kuckuck’s field diaries
(Munda, 2000). During a thorough survey of the west Istrian Coast
in 1950, the same species were found by Ercegović (1952) with
the addition of C. humilis, which was considered a subspecies of C.
compressa at the time. Two of these species are not considered in
the context of this study (C. amentacea var. spicata (Ercegović) G. Fig. 6. A scatterplot with a fitted linear regression line of thallus wet weight (w.w.)
Giaccone and C. crinita Duby) because their distribution is usually against perennial parts w.w. for Cystoseira corniculata in control areas; n = 60.
 M. Devescovi / Aquatic Botany 121 (2015) 26–32
Table 4
Tukey’s HSD tests (P < 0.05) for the interaction S × T × A tested by the ANOVA
reported in Table 3.
Among S levels within combinations of T and A levels
T1, C1: barb < spin < foen < corn T2, C1: barb < spin = corn < foen
T1, C2: barb < foen = spin < corn T2, C2: spin < corn = barb < foen
T1, C3: barb = spin = foen < corn T2, C3: barb = spin = corn < foen
Between T levels within combinations of S and A levels
C1, barb: T2 = T1 C2, barb: T1 < T2 C3, barb: T1 = T2
C1, corn: T2 < T1 C2, corn: T2 < T1 C3, corn: T2 < T1
C1, foen: T1 < T2 C2, foen: T1 < T2 C3, foen: T1 < T2
C1, spin: T2 = T1 C2, spin: T2 < T1 C3, spin: T2 = T1
Among A levels within combinations of T and S levels
T1, barb: C3 = C1 = C2 T2, barb: C1 = C3 < C2
T1, corn: C1 < C3 = C2 T2, corn: C1 < C2 = C3
T1, foen: C1 < C2 = C3 T2, foen: C1 = C2 < C3
T1, spin: C1 = C3 < C2 T2, spin: C2 = C1 < C3
Abbreviations for factors and factor levels are provided in Tables 1 and 3.
During the 1970s and 1980s, Cystoseira spp. stands along the
Istrian Coast had been in a phase of regression likely because of
direct discharge of urban sewage, an increased eutrophication of
the northern Adriatic Sea and sea urchin grazing (Munda, 2000;
Hanel, 2002). The same processes were relevant to a regression of
Cystoseira spp. in the northwestern Mediterranean (Thibaut et al.,
2005). The apparent recovery of Cystoseira spp. along the Istrian
Coast paralleled a marked reduction of eutrophication levels of the
northern Adriatic Sea in the period 2000–2009 (Degobbis et al.,
2000; Djakovac et al., 2012) and a decrease of sea urchin densities,
which started during the 1990s (Hanel, 2002).
Marked differences in total cover (Table 1), ranking of species
composing mixed stands and cover of individual species (Table 4)
were detected among control areas, showing that non-assessed
factors, acting differently from area to area, produced variation
in Cystoseira spp. demography. These factors could include, for
example, storm effects, destructive fishing activities and other
anthropogenic impacts due to agriculture and tourism. Without a
reference condition value (i.e., Cystoseira spp. abundances charac-
teristic for the pristine environment of the northern Adriatic Sea),
it is not possible to quantitatively determine the ecological qual-
ity status class (EC) of sampled areas (WFD; EC, 2000). However,
on the basis of obtained results and previous studies conducted
in other Mediterranean regions (Panayotidis et al., 2004; Thibaut
et al., 2005; Ballesteros et al., 2007; Mangialajo et al., 2008), the
water quality of the Istrian Coast could be assessed descriptively.
No Cystoseira species are expected at sites with a “Low” or “Bad” EC
(Panayotidis et al., 2004; Ballesteros et al., 2007); this type of situa-
tion was not noticed in the study area. In ports, stands were mainly
composed of C. compressa and C. humilis, and the other four species
were present as sparse thalli or entirely absent (Fig. 4). An “Inter-
mediate” EC can therefore be assigned to the sampled port areas,
as it is recognised that C. compressa may withstand moderate levels
of pollution (Thibaut et al., 2005; Mangialajo et al., 2008). In con-
trol areas, all historically recorded disturbance-sensitive Cystoseira
spp. associated with pristine conditions were present, suggesting a
“Good-High” EC.
Overall, the obtained results show that C. barbata, C. corniculata,
C. foeniculacea and C. spinosa are sensitive to levels of disturbance
characteristic for ports of Istrian coastal cities. The species C. bar-
bata, C. foeniculacea and C. spinosa have been considered sensitive in
other Mediterranean regions (Thibaut et al., 2005; Ballesteros et al.,
2007; Sales and Ballesteros, 2009). In contrast however, C. humilis
was found thriving in abundance in Istrian ports and is considered
a sensitive species in the western Mediterranean Sea (Ballesteros
31
et al., 2007). However, it has been suggested that sensitivity levels
for Cystoseira spp. might change according to the Mediterranean
region that is being assessed (Ballesteros et al., 2007).
The sensitivity of northern Adriatic Cystoseira spp. correlated
with their growth patterns. In Cystoseira spp., axis length increases
proportionally with age (Ballesteros et al., 2009). Hence, the thalli
of C. barbata, C. foeniculacea and C. spinosa can be considered much
older than those of C. compressa and C. humilis (Fig. 5). C. corniculata
thalli attained biomass values much higher than those of all other
species. Because the weight of perennial parts (mainly secondary
axes) represented a large proportion of total biomass (Fig. 6), this
species can also be considered to be long-lived. For C. compressa
and C. humilis, assessed axis lengths indicate ages between 1 and
3 years (Ercegović, 1952). In spite of this, due to an extraordinar-
ily rapid growth of primary and higher order branches (deciduous
parts), they attained a thallus biomass similar to that of older C.
barbata, C. foeniculacea and C. spinosa. A rapid growth and likely a
high recruitment rate implied by the high frequency of young thalli
could contribute to the persistence C. compressa and C. humilis in
ports. However, it is also possible that the presence of adaptation
mechanisms lacking in the other species allowed them to dominate
these areas.
The cover of C. compressa and C. humilis did not vary between the
two considered bottom topographic types, and C. compressa was
more abundant in ports than in control areas (Table 2). Previous
studies have documented an increase in C. compressa abundance in
moderately polluted areas (Thibaut et al., 2005; Mangialajo et al.,
2008). It has been proposed that an increased abundance of C.
compressa could be due to diminished competition for space with
other Cystoseira species (Mangialajo et al., 2008), which is in accor-
dance with the Cystoseira total cover decrease detected in Istrian
ports (Table 1). However, the abundance of C. humilis in ports and
control areas was similar. During Cystoseira spp. sampling, it was
observed that C. humilis tended to grow as an epiphyte on other
host Cystoseira spp. or on adventitious calcareous tubes of the lime-
stone boring bivalve Rocellaria dubia (Pennant, 1777) siphons. The
decrease in host Cystoseira spp. in ports could be compensated by
an increased growth on the tubes of R. dubia siphons, thus allowing
an abundance of C. humilis similar to that in control areas.
The results obtained for the other four species suggest that their
variation in abundance could be generated by a combination of
anthropogenic, biological and hydrodynamic processes modulated
by bottom topography. In ports, C. barbata, C. corniculata, C. foenic-
ulacea and C. spinosa regressed; however, they tended to persist
on T1 in higher amounts than on T2 (Fig. 4b). More structured
bottom topography could furnish a refuge against physical pertur-
bations and sediment scouring, which are expected to increase in
ports. However, in sites subjected to nutrient pollution, the graz-
ing control of epiphytic filamentous algae, which overgrow and
negatively affect the underlying community, depends on the com-
position of fish and invertebrate assemblages (Jephson et al., 2008;
Moksnes et al., 2008). The complexity of the substrate topography
positively correlates with total fish abundance, diversity and the
abundance of individual species (McCormick, 1994). Hence, I sug-
gest that increased grazing control of filamentous algae, associated
with the more complex substrate, allowed the survival of the four
species mentioned.
In control areas, C. corniculata and C. foeniculacea were dominant
on bottom types T1 and T2, respectively (Table 4). This pattern could
be related to a difference in the strength by which these two species
are attached to the substrate. During sampling, it was observed that
C. corniculata was very weakly attached to the substrate by forma-
tions originating from creeping secondary axes, which adhered on
the rocky substrate and also to pebbles and shells of dead mol-
luscs. In contrast, C. foenicolacea was very firmly attached to the
rocky substrate by a large basal disc. More erect thin main axes
32 M. Devescovi / Aquatic Botany 121 (2015) 26–32

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