BULLETIN OF MARINE SCIENCE, 78(1): 71–82, 2006

TELEOSTEAN FISH ASSEMBLAGES IN AN

ARTIFICIAL REEF AND A NATURAL ROCKY AREA IN
CATALONIA (NORTHWESTERN MEDITERRANEAN):
AN ECOMORPHOLOGICAL APPROACH

Laura Recasens, Antoni Lombarte, and Pilar Sánchez

Abstract
We used landmark shape analysis to describe the morphospace of fish communi-
ties on an artificial reef and nearby rocky area with a muddy-sand bottom. Twenty-
one monthly fish samples were taken using a commercial trammel net from 2000 to
2003 in the Calafell area (Catalonia, NW Mediterranean). The data were standard-
ized to the number of individuals and weight (g) per 1000 m. In total, 34 teleostean
fish species were used to calculate the ecological parameters of richness, evenness,
and diversity. Geometric morphometry based on landmark analysis revealed that
the communities in the two study areas had a similar morphospace. In both areas
the morphospace was structured into three main groups: benthic fish (flatfish), epi-
benthic species, and nectonic species. The ecomorphological results suggest that
the artificial reef community had a similar species composition and relative abun-
dance to the natural rocky area nearby.

Artificial reefs have shown to be a relatively cheap way of preventing illegal trawl-
ing in Mediterranean littoral ecosystems (Bayle-Sempere et al., 1994; Moreno et al.,
1994). A program to install artificial reefs in Catalonia started in 1978 and continues
to this day. Currently, there are a total of 22 artificial reefs along the coast between
40.5° and 42.5° N. These reefs have mainly been deployed between depths of 15 and
30 m in order to avoid illegal trawling (minimum legal depth for trawlers is 50 m).
Artificial reefs can be important as feeding, spawning, or protection areas for some
fish species. However, due to the concentration of organisms, the structures can put
pressure on species that are more vulnerable to artisanal fishing (Bohnsack, 1989).
In fact, artificial reefs imply a substrate impact because they introduce a rocky sys-
tem into an area where originally there were no rocks. This can produce significant
changes in species composition and assemblages (Coll et al., 1998; Sánchez-Jerez et
al., 2002).
There are different approaches for analyzing communities and determining fish
assemblages. Most methods use specific composition where all species have an in-
dependent weight without considering the species’ ecologic role. Recently, other
aspects such as morphology have begun to be considered, and an ecomorphologic
approach may be a useful new method. Ecomorphology is based on the premise that
the ecology of an organism is related to its morphology (Bock, 1990; Motta et al.,
1995b). One of the main objectives of ecomorphological comparisons is to deter-
mine the relationship between the morphological characteristics and how ecological
resources in a particular environment are used (Karr and James, 1975; Norton et al.,
1995). Given that environments constrain both morphology and ecology, we should
be able to predict ecological patterns for species assemblages from the morphospace
(multidimensional space generated by a multivariable analysis of quantitative mor-
phometric characters; Wiens and Rotenberry, 1980; Motta et al., 1995b). Therefore,

Bulletin of Marine Science 71

© 2006 Rosenstiel School of Marine and Atmospheric Science
of the University of Miami
72 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 1, 2006

the source partitioning (or structure) in a community can be determined by analyz-

ing the morphological characteristics (Ross, 1986).
One of the main problems in ecomorphological studies is deciding which morpho-
logical and ecological traits are the most suitable to use (Norton et al., 1995). These
characteristics should have a close relationship with the environmental factors that
determine the organism’s niche (Bock, 1990; Norton et al., 1995). Among the mor-
phological characteristics that can be analyzed, fish body shape can be related to lo-
comotion efficiency and microhabitat use (Hertel, 1966; Motta et al., 1995a; Schmid
and Senn, 2002).
A problem when using morphological analyses is how to quantify fish shape. The
linear body measurements are easy to use, but these kinds of measurements are re-
lated more to size differences (relative or not) than to shape differences (Lleonart et
al., 2000). The recently developed warp analysis based on comparing morphological
homologous points (landmarks; Rohlf, 1990; Rohlf and Marcus, 1993) allows us to
make a quick, accurate shape measurement. Therefore, landmark analyses can inter-
pret the morphological variability (Lombarte and Recasens, 2004).
Our aim is to describe the morphospace of two fish assemblages on an artificial
reef and a nearby natural rocky area at the same depth, based on their corresponding
morphospace determined by landmark shape analysis (warps). This is complement-
ed by a more classical approach based on ecological descriptors (richness, diversity,
evenness). The assemblage description is based on the specific composition of both
areas.

Materials and Methods

Ecological Descriptors of the Fish Assemblages.—Fish assemblages were examined

at an artificial reef and a nearby rocky area with a muddy-sand bottom at 16–17 m depth.
The Calafell artificial reef is located near the Catalan coast, which is part of the Iberian coast
in the NW Mediterranean (Fig. 1A). This artificial reef consists of a group of 20 production
modules that were installed in 1987 (Alvf1) and another group of 20 production (Alvf2) and
215 mixed modules installed in 1998 (Fig. 1B).
Bi-monthly sampling was carried out in both the artificial reef and the natural rocky zone
during the 2000–2003 time period. Twenty-one fish samples were taken with an artisanal
fishing vessel using a commercial trammel net. Each sample catch was sorted, and all fish col-
lected were identified to the species level, counted, and weighed (g). Data obtained have been
standardized to the number of individuals and weight (g) per 1000 m of trammel net. In total,
34 species were identified (Table 1) and the following ecological parameters were calculated
(Magurran, 1988):
Species richness (Margalef):

S −1
DMg =
ln ( N )

where S is the number of species and N is the number of individuals in the sample; the Shan-
non-Wiener diversity index:

s
H ′ = − ∑ pi ln ( pi )
i =1
 RECASENS ET AL.: FISH ASSEMBLAGES IN ARTIFICIAL AND NATURAL REEFS 73

Figure 1. A) Location of the Calafell artificial reef and natural rocky area sampled in the western
Mediterranean. B) Module types deployed in the Calafell artificial reef.

where pi = proportion of individuals found of the ith species; and equitability, Pielou’s even-
ness:

H′
J=
ln ( S )

where H´ is Shannon-Wiener’s diversity index and S is the number of species.

Morphospace of Fish Assemblages.—The structures of both fish assemblages (artificial
reef and natural rocky areas) were described according to their respective morphospaces ana-
lyzed using geometrical morphometric analysis (Neige, 2003). These analyses were based on
standardized images of the left side of the fish. We analyzed a total of 34 species (28 species in
the artificial reef and 26 species in the rocky area; Table 1). For each species, digitized images
of a maximum of five specimens were obtained. For each specimen 27 landmarks and semi-
landmarks were selected (Fig. 2A). After digitizing the landmark maps, they were rotated,
scaled (to unit centroid size), and translated using a generalized least square superimposition
(GLS) procedure (generalized procrustes) to eliminate scale and orientation distortions (us-
ing tpsRel 1.24; Rohlf, 2001). A thin-plate spline procedure was used to fit an interpolated
function to an average map (consensus configuration) of the carapace shape and derive the
uniform (relative warps) components of shape variation (Fig. 2B). Shape changes were visual-
ized using relative warp analysis (equivalent to the principal components; Rohlf and Marcus,
1993; Adams et al., 2003).
Ten first relative warp scores were selected as variables for describing each consensus spe-
cies shape. In order to determine the degree of shape disparity in the fish assemblages, the
nearest nodal distance between the consensus shape of each species was calculated. This
method is based on applying the mean nearest nodal distance (MND), described by Webb
(2000), to morphological measurements. To determine the Euclidean nodal distances be-
tween species, a cluster analysis was carried out. The Euclidean distance (Sokal and Rohlf,
74 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 1, 2006

Table 1. Species composition, abundance, size, and fish density in the Calafell artificial reef and a natural rocky
area. CV = coefficient of variation. Data on biomass and density are standardized to 1000 m of trammel net.

Artificial reef Rocky area

Species Mean ind. Biomass Biomass Density Mean ind. Biomass Biomass Density
weight (g) mean (g) CV (num) weight (g) mean (g) CV (num)
Boops boops 75.0 6.25 1.97 0.20 77.5 6.46 2.76 0.08
Bothus podas 60.2 58.89 2.54 0.99 62.1 232.52 2.29 3.70
Chelidonichthys lucernus 169.4 72.17 1.68 0.42 - - - -
Chelon labrosus 1,075.0 112.31 1.95 0.11 1,386.3 277.09 1.98 0.18
Chromis chromis 25.0 2.40 2.01 0.20 21.0 2.02 2.83 0.10
Conger conger 632.0 94.80 3.61 0.15 - - - -
Coris julis 35.0 5.25 1.98 0.15 25.0 3.50 2.79 0.14
Dactylopterus volitans - - - - 474.2 68.79 2.04 0.13
Diplodus annularis 63.4 9.07 2.28 0.14 79.5 23.98 2.23 0.28
Diplodus sargus 302.8 158.80 1.40 0.53 328.7 329.38 1.09 0.98
Diplodus vulgaris 231.0 35.46 2.20 0.18 - - - -
Lithognatus mormyrus 424.4 100.31 2.01 0.23 372.8 256.31 2.83 0.69
Merluccius merluccius 245.8 56.73 3.61 0.23 - - - -
Mullus barbatus 65.0 8.59 2.03 0.13 77.0 22.81 1.92 0.30
Mullus surmuletus 137.0 57.64 2.98 0.34 149.3 54.11 2.48 0.35
Pagellus acarne 64.0 144.18 0.99 2.25 80.5 204.31 0.65 2.61
Pagellus erythrinus 198.3 666.03 1.90 3.18 263.6 527.94 1.03 1.98
Pagrus pagrus 291.8 36.15 3.44 0.19 - - - -
Phycis phycis 317.3 241.64 1.37 0.59 201.2 62.77 1.23 0.31
Pomadasys incisus 90.3 21.32 2.05 0.23 - - - -
Psetta maxima 191.0 29.39 3.61 0.15 - - - -
Sciaena umbra 691.3 59.08 2.66 0.09 597.0 300.58 1.86 0.50
Scophtalmus rhombus 163.3 141.68 2.41 0.87 126.6 181.83 1.81 1.44
Scorpaena notata 86.4 895.77 1.86 10.54 72.7 173.29 1.41 2.52
Scorpaena porcus 99.8 96.97 1.59 0.69 129.0 159.95 2.17 1.26
Scorpaena scrofa 218.1 43.90 1.67 0.21 413.3 311.86 1.29 0.78
Seriola dumerilii 555.0 127.24 2.11 0.22 - - - -
Serranus cabrilla 101.4 39.76 1.40 0.44 100.2 60.75 1.16 0.64
Solea senegalensis 342.4 375.54 1.37 1.08 286.4 178.30 1.56 0.58
Solea solea 179.2 130.51 1.57 0.68 249.1 162.88 1.75 0.73
Symphodus tinca - - - - 297.0 70.87 1.91 0.25
Trachurus mediterraneus - - - - 236.9 51.20 2.63 0.43
Umbrina canariensis - - - - 331.3 127.40 2.83 0.38
Uranoscopus scaber - - - - 229.0 38.17 1.87 0.17

1962) and the UPGMA aggregation algorithm (Sokal and Michener, 1958) were employed
because these methods yield a high cophenetic correlation coefficient.

Results

Specific Composition.—Specific composition, size, biomass, and density are

shown in Table 1. The species that comprise the highest proportion of the biomass in
the artificial reef area were Scorpaena notata Rafinesque, 1810, Pagellus erythrinus
(Linnaeus, 1758), and Solea senegalensis Kaup, 1858, whereas in the rocky area P.
 RECASENS ET AL.: FISH ASSEMBLAGES IN ARTIFICIAL AND NATURAL REEFS 75

Figure 2. A) Location of the 27 landmarks on the left lateral side of the fish’s body used in the
geometric morphologic analyses (warps). B) Thin-plate spline from the average map (consensus
configuration) of Diplodus sargus specimens.
erythrinus, Diplodus sargus (Linnaeus, 1758), and Scorpaena scrofa Linnaeus, 1758
were the most dominant species. The CV (coefficient of variation: standard devia-
tion/mean) of biomass tended to be slightly lower in the natural rocky area than in
the artificial reef. The species with lowest biomass CV were Pagellus acarne (Risso,
1827), Phycis phycis (Linnaeus, 1766), and S. senegalensis in the artificial reef, and
P. acarne, P. erythrinus, and Serranus cabrilla (Linnaeus, 1758) in the rocky area.
In terms of abundance, S. notata and P. acarne were abundant in both areas and
S. notata was the most abundant species in the artificial reef. Pagellus erythrinus
was more common in the artificial reef area and Bothus podas (Delaroche, 1809)
were more common in the rocky area. Some species were absent in one of the areas,
such as Dactylopterus volitans (Linnaeus, 1758), Simphodus tinca (Linnaeus, 1758),
Trachurus mediterraneus (Steindachner, 1868), Umbrina canariensis Valenciennes,
1843, and Uranoscopus scaber Linnaeus, 1758, which were absent in the artificial
reef, and Chelidonichtys lucernus (Linnaeus, 1758), Conger conger (Linnaeus, 1758),
Merluccius merluccius (Linnaeus, 1758), Seriola dumerilii (Risso, 1810) among oth-
ers, which were not found in the rocky area.
Species richness, evenness, and diversity tended to be higher in the rocky area than
in the artificial reef area (Table 2), but these differences were not significant except
for richness.
Morphological Structure.—The morphospace obtained from the geometrical
morphology analysis (warp analysis) had a similar structure in both areas studied
76 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 1, 2006

Table 2. Ecological descriptors (richness, evenness, and diversity) of fish assemblages in the
Calafell artificial reef and natural rocky area. t-test used for comparison among areas (P < 0.05).

Richness Evenness Diversity

Artificial reef
Mean 2.10 0.73 1.24
SD 0.55 0.20 0.31
Rocky area
Mean 3.40 0.84 1.50
SD 1.48 0.12 0.33

(Figs. 3A, 4A). In the three first warp representations of the morphospace, the con-
sensus configuration of each species is represented by one point. The three first warps
of the fish assemblage in the artificial reef area represent 80.69% of the total morpho-
logical variability (Fig. 3A). In the natural rocky area the accumulated morphological
variability was 76.47% (Fig. 4A). In both analyses, the first warp is the proportion of
cephalic size in relation to body length. Thus, the negative values represent species
with relatively small head sizes (flatfishes and C. conger) and the most positive values
species with relatively large heads (Scorpaenidae). There is also a central group with
intermediate characteristics that is more evident in the natural rocky area, than in
the artificial reef where this group is mixed with the positive value group. The second
warp expresses the relationship between body length and body width. Elongated fish
such as C. conger have negative values in the second warp, and rounded fish such as
B. podas or Diplodus spp. have positive values.
In the artificial reef analyses, the third warp is related to the anatomy of the anal
fin. Negative values indicate a rounded fin, such as in C. conger, and positive values
represent a forked shape, such as in Chelon labrosus (Risso, 1827), Mullus spp., or
Chromis chromis (Linnaeus, 1758). In the rocky area, the most negative value of the
third warp is associated with the extraordinary development of the pectoral fin of D.
volitans (a species that is not present in the artificial reef area). The third warp’s most
positive values are represented by C. labrosus, which is characterized by a relatively
small pectoral fin.
Morphological Diversity.—The cluster analysis (Euclidean distance and UPG-
MA) calculated from the ten first warp scores (more than 96% of the total accumulat-
ed morphological variability in both warp analyses) also showed a similar structure
in both study areas with more structured groups in the natural rocky area; flatfishes
were the most clearly differentiated group (Figs. 3B, 4B). In the species groups ob-
tained from the cluster analysis for both areas, the maximum distance, chaining, and
cophenetic correlation coefficient were similar (Table 3). The mean nearest nodal dis-
tance (MND) as an expression of the morphological diversity falls within the same
range in both fish assemblages (Table 3), although in the rocky area the MND value
(0.156) expressed larger morphological diversity than the MND value of the artificial
reef assemblage (0.147).

Discussion

The effect artificial reefs have on the areas in which they are installed has long
been an issue of debate. Debates have centered on the controversy that considers
artificial structures to be attractors and redistributors of organisms that previously
 RECASENS ET AL.: FISH ASSEMBLAGES IN ARTIFICIAL AND NATURAL REEFS 77

Figure 3. A) Representation of the three first warps based on the 27 landmarks of the Teleostean
species in the artificial reef. The size of the point represents the mean weight of the individual,
and the color of the point represents the density (standardized number of specimens per sample).
B) Cluster analysis (Euclidean distance, UPGMA) based on the 10 first warp scores of the artifi-
cial reef assemblages. * Indicates species not present in the natural rocky area.
inhabited these areas (Bohnsack, 1989; Prat, 1994), versus structures with a high
biomass production that contributes to regenerating degraded areas and increasing
population abundance (Ambrose and Swarbrick, 1989; Stephens and Pondella, 2002).
These points of contention have been difficult to resolve.
In Iberian Mediterranean areas installing these artificial structures has been
aimed at preventing illegal trawling in coastal zones (Recasens et al., 2001). Tradi-
tionally, coastal zones have been exploited by an artisanal fleet that captures large
individuals with a high commercial value (Farrugio and Le Corre, 1993). Due to the
Mediterranean’s physical and biological characteristics, such as being an oligotro-
phic, low productive sea (Estrada et al., 1985), large increases in fish production as
a result of introducing artificial reefs are not expected. However, it is interesting to
determine the organization level of these communities. Comparison with a nearby
78 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 1, 2006

Figure 4. A) Representation of the three first warps based on the 27 landmarks of the Teleostean
species in the natural rocky area. The size of the points represents the mean weight of the indi-
vidual, and the color of the point represents the density (standardized number of specimens per
sample). B) Cluster analysis (Euclidean distance, UPGMA) based on the 10 first warp scores of
the natural rock assemblages. * Indicates species not present in the Calafell artificial reef.

natural rocky area has allowed us to show that at an ecological descriptor level, ar-
tificial reefs tend to be similar to natural rocky areas, but they have lower values in
these ecological descriptors especially in richness. This could indicate a lower level
of assemblage organization that could be related to differences in habitat structure
at small-scale (García-Charton et al., 2004). The trends obtained in this study are
comparable to other studies (Carr and Hixon, 1997).
In terms of the morphological characterization of the fish assemblages, the mor-
phospace structures of the artificial reef and natural rocky areas are similar. In both
cases the two first warps, which represent more than 75% and 68%, respectively, of
the morphological variability in the fish assemblages, are related to the same char-
acteristics. The first warp expresses the differences in head proportion in relation
to body length. This proportion is directly related to the fish’s locomotion systems
 RECASENS ET AL.: FISH ASSEMBLAGES IN ARTIFICIAL AND NATURAL REEFS 79

Table 3. Mean nearest nodal distances (MND) and their standard deviations (SD), cophenetic
correlation coefficient (CCC), chaining (C), and maximum distance (MD) of the cluster analysis
based on the warp scores of geometrical morphologic analyses of the Calafell artificial reef and
natural rocky area.

Artificial reef Natural rocky area

MND 0.147 0.156
SD 0.101 0.092
CCC 0.930 0.917
C 0.291 0.353
MD 0.430 0.434

(Hertel, 1966; Lindsey, 1978). Pleuronectiformes and Anguilliformes with a relatively

small head length (negative first warp values) are characterized by Anguiliform mode
locomotion involving the entire body. The central values represent species such as
Gadiformes and Carangidae that typically use the posterior portion of the body to
move (Subcarangiform and Carangiform mode). The species with a relatively larger
head size have a Labriform mode of locomotion based on oscillating their pectoral
fins (Lindsey, 1978). The fact that these three groups are more evident in the natural
rocky area than in the artificial reef could reveal a more structured community in
the natural system. By combining the three first warps, the species are distributed
in relation to their habitat preferences as observed by Motta et al. (1995a). Benthic
species, such as flatfish are located on one side of the morphospace. These fish prefer
muddy, sandy sea-bottoms (Massutí and Reñones, 2005). On the opposite side of the
morphospace we find the epibenthic or benthic species that live around and over the
natural rocks or artificial modules, such as Sparidae, Sciaenidae, Labridae (Sympho-
dus), Serranidae, and Scorpaenidae (García-Rubies, 1997). Finally, in the central zone
of the morphospace we find the pelagic (Carangidae, Boops boops (Linnaeus, 1758))
and epibenthic swimmer species (Gadiformes) (Lighthill, 1969; Lindsey, 1978). When
comparing morphologically related species that are found in the same morphospace,
other aspects such as differences in size and density can express the ecological seg-
regation that minimizes competition (Schoener, 1974; Ross, 1986; Labropoulu and
Eleftheriou, 1997; Lombarte et al., 2000), as observed in our results. That is, mor-
phologically closely related species pairs, such as D. sargus and Diplodus annularis
(Linnaeus, 1758), S. scrofa and Scorpaena porcus Linnaeus, 1758, and P. erythrinus
and P. acarne, differ in size and abundance. On the other hand, differences in the
specific composition in both locations (natural and artificial) could also be related
to the ecological niche the species occupies. Therefore, there are different species
that are present in one area and not in the other, but which occupy a similar ecologi-
cal niche. This could be the case for S. dumerilii, which is present in the artificial
reef, and T. mediterraneus, present in the natural rocks, both of which are swimmer
species, and also C. lucernus and D. volitans, in this case benthic species with big
pectoral fins. The presence/absence of some other species such as C. conger, M. mer-
luccius, and Psetta maxima (Linnaeus, 1758) seems to be related to their general low
abundance in the study areas, as these species had a high CV of biomass. However,
the most regular species in both areas is P. acarne, which presents the lowest CV of
biomass. Quantifying the shape disparity not only reflects the habitat partitioning
in the fish assemblage, but also permits us to make a complementary description of
their diversity (Neige, 2003), based on calculating the mean nearest nodal distance
80 BULLETIN OF MARINE SCIENCE, VOL. 78, NO. 1, 2006

(MND). This distance was used to describe diversity in quantitative data related to
the phylogeny of forest tree assemblages (Webb, 2000) and microbial communities
(Bohannan and Hughes, 2003). The morphological diversity obtained from the MND
can be used to complement the classic ecological indexes of richness and evenness
because these indexes include the degree of relationship (in this case based on their
phenotype) between the species that form the assemblage (Bohannan and Hughes,
2003). Therefore, our results can be compared with other ecological indices. Similar
to these indices, morphological diversity reveals that the artificial reef is similar to
the natural rocky area, but that it has generally lower values. Thus, morphological
warp analysis is a fast and adequate tool for describing morphological diversity and
structure based on source partitioning of a fish community.

Acknowledgements

The authors wish to thank M. Demestre, D. Díaz, D. Lloris, G. Fuster, and the ESCAL proj-
ect team for their help during the sampling period. We also acknowledge the skill of the
fisherman J. R. Nuñez and the Fishermen’s Association of Calafell. This work was supported
by the Catalan Government (General Management of Fishing and Maritime Matters). Com-
ments of I. Moreno and A. Gordoa improved the quality of the manuscript.

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Address: Institut de Ciències del Mar (ICM-CSIC). Passeig Marítim 37-49, 08003 Barcelona
Spain. Corresponding Author: (L.R.) E-mail: <laura@icm.csic.es>.