Reviews in Fish Biology and Fisheries 11: 217–254, 2002.

© 2002 Kluwer Academic Publishers. Printed in the Netherlands.

217

Feeding habits and trophic levels of Mediterranean fish

Konstantinos I. Stergiou1 & Vasiliki S. Karpouzi1,2

1 Aristotle
University of Thessaloniki, School of Biology, Department of Zoology, Laboratory of Ichthyology, Box
134, 54124 Thessaloniki, Hellas (Greece) (E-mail: kstergio@bio.auth.gr; Phone: 30310-998268; Fax: 30310-
998279); 2 Present address: University of British Columbia, Fisheries Centre, 2204 Main Mall, Vancouver B.C.
V6T 1Z4, Canada

Accepted 7 May 2002

Contents
Abstract page 217
Introduction 218
Materials and methods 219
Results 234
Discussion 239
Acknowledgements 248
References 248

Key words: feeding habits, fish, fractional trophic levels, Mediterranean Sea

Abstract

The estimation of fractional trophic levels (TROPHs) is essential for the management of fisheries resources
as well as for quantifying the ecosystem effects of fishing. We gathered all available information concerning the
feeding habits of 332 fish stocks, belonging to 146 species, 59 families and 21 orders, throughout the Mediterranean
Sea, and estimated their TROPH values. The latter ranged from 2.0 to 4.5 and the following functional trophic
groups were identified: (a) pure herbivores (TROPH = 2.0–2.1, mean = 2.02, SD = 0.03), which were very rare and
represented by Siganus luridus, Siganus rivulatus and Sarpa salpa, all of which feed on red, brown, green and blue-
green algae; (b) omnivores with a preference for vegetable material (2.1 < TROPH < 2.9, mean = 2.5, SD = 0.12),
but feeding on other prey, such as sponges, hydrozoans, anthozoans, polychaetes, ostracods, isopods, amphipods
and copepods. This type of omnivore was very rare among the cases reviewed; (c) omnivores with a preference
for animal material (2.9 < TROPH < 3.7, mean = 3.4, SD = 0.19) feeding on a wide variety of prey (e.g., algae,
foraminifera, brachyurans, balanoids, ascidians, amphipods, appendicularians, annelids, isopods, gastropods,
cnidarians, ophiurids, polychaetes, cladocerans, mysids, euphausids, fish larvae, cephalopods). They were the
most numerous and were mainly represented by species of the families Blenniidae, Bothidae, Centracanthidae,
Gobiidae, Labridae, Lotidae, Macrouridae, Mullidae, Ophidiidae, Soleidae, Triglidae and Engraulidae; and (d)
carnivores with a preference for large decapods, cephalopods and fish (3.7 < TROPH < 4.5). They were the
next most abundant group among the cases reviewed. They were mainly represented by species of the families
Dalatiidae, Lophiidae, Scombridae, Scyliorhinidae, Synodontidae, Torpedinidae, Merlucciidae, Xiphiidae and
Zeidae. This group was divided into two subgroups: one exhibiting a preference for decapods and fish (3.7 <
TROPH < 4.0, mean = 3.85, SD = 0.09) and another one exhibiting a preference for fish and cephalopods (4.0 <
TROPH < 4.5, mean = 4.38, SD = 0.12). Potential top carnivores in the Mediterranean Sea were also identified.
Although such a grouping is provisional, and subject to revision when more datasets for other species, habitats
and areas (especially from the Southern Mediterranean Sea, which was underrepresented in our compilation)
218

become available, it may serve as a basis for the maintenance of trophic level balance in the Mediterranean Sea.
This will set the basis for an ecosystem approach to the management of the Mediterranean Sea. Across species,
TROPH values increased asymptotically with maximum reported body length. The same was also true for 9 out
of 11 stocks, for which diet composition data were available for more than three length classes. The within- and
among-species asymptotic relationships between TROPH and body length are discussed within the framework of
foraging behavior and prey selection theory. Finally, based on the sampling characteristics and methodology used
in the studies reviewed, some recommendations for future stomach content studies are also provided.

Introduction
The quality and quantity of food are among the
most important exogenous factors directly affecting
growth and, indirectly, maturation and mortality in
fish, thus being ultimately related to fitness (Wootton,
1990). Traditionally, information on the quality and
quantity of food consumed by fish, which can be
derived from feeding studies, is made operational for
fisheries research through incorporation into appro-
priate fisheries models (e.g., multispecies virtual
population analysis) and, after scaling up to the
total biomass of predators and prey, provides esti-
mates of the total biomass consumed by predators
(Jennings et al., 2001). This allows: (a) the assess-
ment of the relative importance of predatory mortality
when compared to fishing mortality for commercially
exploited populations; (b) the exploration of any links
between recruitment variability and predation (Link
and Almeida, 2000); and (c) variability in feeding to
be effectively used for the study of the dynamics of
fish populations and thus to increase the accuracy of
resulted predictions (Ulltang, 1996).
In addition, diet composition data also play a
key role for the research on the following ecolog-
ical issues: (a) resource partitioning and within-
and between-species competition (e.g., Macpherson,
1981; Harmelin-Vivien et al., 1989); (b) prey selection
(e.g., Kohler and Ney, 1982; Stergiou and Fourtouni,
1991); (c) predator-prey size relationships (e.g., Pauly,
2000a; Scharf et al., 2000); (d) distribution of feeding
types with latitude (e.g., herbivory versus latitude;
Pauly, 2000b); (e) ontogenetic diet shifts (e.g.,
Stergiou and Fourtouni, 1991; Labropoulou et al.,
1997); (f) habitat selection (e.g., Labropoulou and
Machias, 1998; Labropoulou et al., 1999); (g) testing
predictions from foraging behaviour and optimal
foraging theory models (e.g., Wootton, 1990; Hughes,
1997); and (h) species invasions (e.g., Stergiou, 1988;
Golani and Galil, 1991).
Finally, diet composition data are also used for the
estimation of trophic levels (Pauly and Christensen,
2000a; Pauly and Sa-a, 2000), the latter being of
paramount importance for the management of aquatic
resources. Recent, paradigmatic failures in fisheries
management clearly indicate that traditional assess-
ment models, management strategies, and reference
points derived from them are largely inadequate to
satisfy the purpose of their existence (e.g., De La
Mare, 1998; Beverton, 1998; Smith, 1998; Stokes
et al., 1999; Stergiou, 2002). Such a failure is real-
ized from the highly diversified and complex effects of
overfishing on the life histories of individual species
and on the ecosystems in which these species are
embedded (e.g., Jennings and Kaiser, 1998; Hall,
1999; Jennings et al., 2001; Stergiou, 2002). For
instance, the mean trophic level in fisheries landings
(i.e., mean trophic level of all species making up the
landings weighted by their catch) in the last 45 years
has decreased steadily both at the global scale and
at the regional, ocean-specific scale (e.g., Atlantic,
Pacific, Mediterranean; the “fishing down the food
web” effect: Pauly et al., 1998a, b, 2000a). The failure
of traditional management approaches has led to a
growing interest in “ecosystem-based management”
as a promising alternative strategy (e.g., Jennings and
Polunin, 1996; Walters et al., 1997; Pitcher et al.,
1998; Mooney, 1998; Pauly, 1998a; Gislason et al.,
2000; Cochrane, 2000; Pitcher, 2000; Stergiou, 2002).
It is generally difficult to envisage the full implications
of ecosystem management, mainly because the inter-
actions involved at the ecosystem level are complex
and poorly understood (Jennings et al., 2001). As a
result, it is also difficult to develop a full array of
clearly defined “ecosystem” objectives, indicators and
reference points that will trigger management actions,
without which ecosystem management cannot be real-
ized. Yet, there is a growing understanding that trophic
level balance might serve as an objective (Pauly et
al., 1998a, b, 2000a; Briand, 2000; Gislason et al.,

2000), which can be coined to clearly defined indi-
cators (e.g., “fishery is balanced”, FIB, index (Pauly
et al., 2000b), aggregate annual removals from each
trophic level) and reference points (e.g., maximum
value of FIB index; maximum % annual removals,
respectively) (see Table 1 in Gislason et al., 2000).
Furthermore, the estimation of trophic levels is
also very useful for quantifying the effects of fishing
on marine ecosystems because it allows the devel-
opment of new approaches to the analysis of marine
food webs such as: (a) accurate estimation of the
“Primary Production Required” to maintain fisheries
(Pauly and Christensen, 1995; Tudela, 2000); (b)
the construction of a series of mean trophic level
values of fish and invertebrates landed by fisheries as
a means of evaluating the impacts of fishing on marine
ecosystems (Pauly et al., 1998a,b, 2000b, 2001;
Christensen, 1998; Stergiou and Koulouris, 2000);
and (c) comparative community analysis, in which
available community studies can be re-expressed using
trophic levels as a common currency (Pauly et al.,
2000c).
Despite the key role of trophic levels for fisheries
and ecosystem research, it is only since the devel-
opment of Ecopath in the early 1980s (Polovina,
1984; Christensen and Pauly, 1990, 1991, 1992a,
b, 1993) and its evolution in the following years
into a highly valuable ecological modeling tool
(Ecopath/Ecosim/Ecospace: Walters et al., 1997,
1999, 2000; Jarre-Teichmann, 1998; Okey and Pauly,
1999; Christensen et al., 2000; Pauly et al., 2000c;
Pauly and Christensen, 2000b; Watson et al., 2000)
and the development of FishBase (Froese and Pauly,
1990, 1994, 1998, 2000; Pauly and Froese, 1992)
that trophic levels gained high importance leading
to global studies (e.g., Christensen and Pauly, 1995;
Pauly and Christensen, 1995; Pauly, 1998b; Pauly
et al., 1998a,b, 2000a) in which previously reported
pieces of information were transformed into thorough
scientific knowledge.
Nowadays the compilation of existing stomach
content data for various aquatic organisms seems to
be one among many necessary steps for the develop-
ment of ecosystem models through the use of various
modeling tools, Ecopath/Ecosim/Ecospace included.
Such models will mediate the development and evalu-
ation of trophic and other reference points (see Table
1 in Gislason et al., 2000) as well as the evaluation
of the effects of fishing on marine ecosystems, by
testing “what if” scenarios of alternative management
policies. Within this general framework, Pauly et al.
219
(1995, 1998c) compiled all available diet composition
data and estimated fractional trophic levels (TROPHs)
for 97 species of marine mammals. This inspired
Cortés (1999) to do a similar compilation for 149 shark
species.
In this review, we gathered all the available studies
concerning the feeding habits of Mediterranean fish
and estimated their TROPHs. Overall, diet composi-
tion data were gathered for 332 datasets, corre-
sponding to 146 species, 59 families and 21 orders.
This information will be useful for an ecosystem-
based management of the Mediterranean Sea. The
inadequacy of “traditional” single-species models is
amplified by the lack of long-term routine fisheries
data and the poor level of scientific information, in
contrast to the considerable increase in fishing activity
over time (Stergiou et al., 1997; Stergiou, 2000;
Lleonart, 2000; Briand, 2000).
Materials and methods
We gathered all available information pertinent to
feeding habits of Mediterranean fish, derived from
stomach content studies conducted since 1961, using:
(a) Aquatic Sciences and Fisheries Abstracts (ASFA);
and (b) the bibliographic database of the Commission
Internationale pour l’Exploration Scientifique de la
Mer Méditerranée (CIESM). Both sources cover peer-
reviewed as well as grey literature articles. We also
used some unpublished theses and technical reports
that were available to us.
All available quantitative and qualitative infor-
mation provided in the original studies was tabu-
lated by species, study area and year, henceforth
called “dataset” (Table 1A). In addition, the following
information was compiled: sampling gear, sampling
frequency, number of stomachs analyzed, method used
for diet expression (Table 1A), range in length or
mean length of specimens studied, main prey species
or group of species by predator’s length when avail-
able, and prey’s quantitative contribution to the diet
(Table 1B). Finally, the maximum reported body
length (Lmax ) and the habitat type of all species
reviewed here were extracted from FishBase online
(www.fishbase.org) (Table 1B).
TROPHs were calculated for each dataset based on
the full array of prey items in the diet as presented
in the original studies. TROPH expresses the posi-
tion of organisms within the food webs that largely
define aquatic ecosystems (Pauly and Christensen,
220
Table 1A. Feeding habits of Mediterranean marine fish. SM: sampling method (T = trawl, H = hooks, N = nets, GN = gill nets, TN =
trammel nets, PS = purse seiners, SC = SCUBA, Sp. = Speared, CL = commercial landings); SF: sampling frequency (S = seasonal, M
= monthly, 1 = once, 2 = twice); N: number of stomachs examined; and Method: parameters estimated (F = frequency of occurrence, N
= numerical percentage, W = percentage by weight, ES = percentage of empty stomachs, RA = relative abundance). † Valid names of
species (from “FishBase online”; www.fishbase.org)

Species Family Area Year SM SF N Method Reference

Anguilliformes
1. Conger conger Congridae Balearic Sea 1976–1978 T M 242 F, W Macpherson (1979a, 1981)
2. Conger conger Congridae G. Lions June 1981 T 1 12 F, N, W, ES Khoury (1984)
3. Conger conger Congridae G. Marseille 1980 T M 6 F, N, W, ES Bell and Harmelin-Vivien (1983)
4. Gnathophis mystax Congridae Balearic Sea 1976-78 T M 218 F, W Macpherson (1979a, c, 1981)
5. Gnathophis mystax Congridae Balearic Sea 1985–1988 – M 707 F, N, W, ES Casadevall and Matallanas (1990)
6. Ophichthus rufus Ophichthidae Catalan Sea 1985–1987 T M 689 F, N, W, ES Casadevall et al. (1994)
Atheriniformes
7. Atherina boyeri Atherinidae W Sicily 1991 T 1 – F Mirto et al. (1994)
Aulopiformes
8. Bathypterois mediterraneus Ipnopidae Balearic Sea July 1987 – 1 50 F, N, ES Carrassón and Matallanas (1990)
9. Saurida undosquamis Synodontidae SE Mediterranean 1980–1981 – M 5223 N Bingel and Avsar (1988)
10. Saurida undosquamis Synodontidae SE Mediterranean – T M 333 N, ES Golani (1993)
11. Synodus saurus Synodontidae SE Mediterranean – T M 57 N, ES Golani (1993)
Beryciformes
12. Sargocentron rubrum Holocentridae SE Mediterranean 1979–1980 TN M 282 F, N, W, ES Golani et al. (1983)
13. Hoplostethus mediterraneus Trachichthyidae Ionian Sea 1996–1997 T M 482 F, N, ES Madurell and Labropoulou (2000)
Carcharhiniformes
14. Galeus melastomus Scyliorhinidae Balearic Sea 1976–1977 T M 1559 F, W Macpherson (1979a, 1980a)
15. Galeus melastomus Scyliorhinidae Balearic Sea 1976–1978 T M 1559 F, W Macpherson (1981)
16. Galeus melastomus Scyliorhinidae Catalan Sea 1988–1990 T – 99 F, N, W, ES Carrassón et al. (1992)
17. Scyliorhinus canicula Scyliorhinidae Balearic Sea 1976–1978 T M 1009 F, W Macpherson (1979a, 1981)
Chimaeriformes
18. Chimaera monstrosa Chimaeridae Balearic Sea 1976–1978 T, H M 206 F, W Macpherson (1979a, 1980b, 1981)
Clupeiformes
19. Sardina pilchardus Clupeidae G. S Evvoikos 1992–1993 GN, TN S 35 F, ES Petrakis et al. (1993)
20. Sardina pilchardus Clupeidae Dardanelles 1959–60 PS S 131 N, ES Demirhindi (1961)
21. Engraulis encrasicolus Engraulidae Catalan Sea 1994–1995 T 2 311 RA Tudela and Palomera (1995, 1997)
22. Engraulis encrasicolus Engraulidae G. Lions 1995 T M 279 F, W, ES Plounevez and Champalbert (2000)
23. Engraulis encrasicolus Engraulidae G. Lions 1996 T M 247 F, W, ES Plounevez and Champalbert (2000)
Gadiformes
24. Gadiculus argenteus argenteus Gadidae Balearic Sea 1976–1977 T M 1434 F, W Macpherson (1978a, c)
25. Gadiculus argenteus argenteus Gadidae Balearic Sea 1976–1978 T M 1434 F, W Macpherson (1981)
26. Micromesistius poutassou Gadidae Balearic Sea 1976–1977 T M 1761 F, W Macpherson (1978a, c)
27. Micromesistius poutassou Gadidae Balearic Sea 1976–1978 T M 1761 F, W Macpherson (1981)
28. Micromesistius poutassou Gadidae G. Patraikos 1984–1985 T S 310 F, N, W, ES Petrakis and Stergiou (1987)
29. Micromesistius poutassou Gadidae G. Korinthiakos 1984–1985 T S 310 F, N, W, ES Petrakis and Stergiou (1987)
30. Micromesistius poutassou Gadidae G. Evvoikos 1986–1987 T S 641 F, N, W, ES Papaconstantinou et al. (1989)
31. Trisopterus minutus† Gadidae N Tyrrhenian Sea 1985–1987 T S 200 F, N, W, ES Biagi et al. (1992)
32. Trisopterus minutus† Gadidae C Adriatic Sea 1985–86 T S 1186 F, N, W, ES Gramitto (1999)
33. Trisopterus minutus† Gadidae G. Evvoikos 1986–1987 T S 590 F, N, W, ES Politou and Papaconstantinou (1994)
34. Trisopterus minutus† Gadidae G. Pagassitikos 1986–1987 T S 482 N, W, ES Politou et al. (1988)
35. Trisopterus minutus† Gadidae N Aegean Sea 1990–1992 T S 296 F, N Papaconstantinou et al. (1993)
36. Gaidropsarus biscayensis† Lotidae Balearic Sea 1976–1978 T M 395 F, W, ES Macpherson (1978b, c, 1981)
37. Gaidropsarus biscayensis† Lotidae C Adriatic Sea 1982–1983 T S 582 F, N, W, ES Gramitto (1985)
38. Gaidropsarus mediterraneus Lotidae G. Lions June 1981 T 1 15 F, N, W, ES Khoury (1984)
39. Gaidropsarus mediterraneus Lotidae G. Marseille 1980 T M 16 F, N, W, ES Bell and Harmelin-Vivien (1983)
40. Gaidropsarus vulgaris Lotidae G. Lions June 1981 T 1 23 F, N, W, ES Khoury (1984)
41. Gaidropsarus vulgaris Lotidae G. Marseille 1980 T M 7 F, N, W, ES Bell and Harmelin-Vivien (1983)
42. Molva macrophthalma† Lotidae Balearic Sea 1976–1978 T M 355 F, W Macpherson (1979a, d, 1981)
43. Caelorinchus caelorhincus† Macrouridae Balearic Sea 1976–1978 T M 160 W Macpherson (1979a, b)
44. Caelorinchus caelorhincus† Macrouridae Balearic Sea 1976–1978 T M 160 F, W Macpherson (1981)
45. Hymenocephalus italicus Macrouridae Balearic Sea 1976–1978 T M 323 W Macpherson (1979a, b)
46. Hymenocephalus italicus Macrouridae Balearic Sea 1976–1978 T M 323 F, W Macpherson (1981)
47. Nezumia aequalis Macrouridae Balearic Sea 1976–1978 T M 168 W Macpherson (1979a, b)
48. Nezumia aequalis Macrouridae Balearic Sea 1976–1978 T M 168 F, W Macpherson (1981)
49. Trachyrincus scabrus† Macrouridae Balearic Sea 1976–1978 T M 1670 W Macpherson (1979a, b)
50. Trachyrincus scabrus† Macrouridae Balearic Sea 1976–1978 T M 1670 F, W Macpherson (1981)
51. Trachyrincus scabrus† Macrouridae G. Genoa 1977–1978 T S 100 N, ES Relini Orsi and Wurtz (1979)
52. Merluccius merluccius Merlucciidae Balearic Sea 1976–1978 T – 664 F, W Macpherson (1979a, 1981)
53. Merluccius merluccius Merlucciidae G. Lions 1993 T M 1526 F, N, W, ES Bozzano et al. (1997)
54. Merluccius merluccius Merlucciidae C Mediterranean 1982–1983 T 2 660 F, N, W, ES Andaloro et al. (1985)
55. Merluccius merluccius Merlucciidae C Adriatic Sea 1963–1964 T M 363 N, ES Jukić (1972)
 221
Table 1A. Continued

Species Family Area Year SM SF N Method Reference

56. Merluccius merluccius Merlucciidae C Adriatic Sea 1971–1973 – S 1217 N Froglia (1973)
57. Merluccius merluccius Merlucciidae C Adriatic Sea – – – 295 F Jardas (1976)
58. Merluccius merluccius Merlucciidae G. Saronikos 1975–1976 T S 451 F, N, W, ES Caragitsou and Tsimenides (1977)
59. Merluccius merluccius Merlucciidae G. Korinthiakos 1983–1984 T S 216 F, N, W, ES Papaconstantinou and Caragitsou (1987a)
60. Merluccius merluccius Merlucciidae Ionian Sea 1983–1984 T S 342 F, N, W, ES Papaconstantinou and Caragitsou (1987a)
61. Merluccius merluccius Merlucciidae G. Patraikos 1983–1984 T S 731 F, N, W, ES Papaconstantinou and Caragitsou (1987a)
62. Merluccius merluccius Merlucciidae Cyclades Islands – – – – RA Kyrtatos (1982)
63. Merluccius merluccius Merlucciidae W Aegean Sea 1990–1991 T S 316 F, N, W, ES Papaconstantinou et al. (1993)
64. Merluccius merluccius Merlucciidae Cretan waters 1990–1992 T M – W Labropoulou and Markakis (1998)
65. Lepidion lepidion Moridae Balearic Sea July 1987 – 1 36 F, N, ES Carrassón and Matallanas (1990)
66. Lepidion lepidion Moridae Catalan Sea 1988–1990 T 2 617 F, N, W, ES Carrassón et al. (1997)
67. Phycis blennoides Phycidae Balearic Sea 1976–1978 T M 2251 F, W, ES Macpherson (1978b, c, 1981)
68. Phycis phycis Phycidae Dodecanese 1985–1986 TN S 196 F, N, W, ES Papaconstantinou and Caragitsou (1989)
Gobiesociformes
69. Apletodon dentatus Gobiesocidae G. Marseille 1980 T M 1 F, N, W, ES Bell and Harmelin–Vivien (1983)
Lophiiformes
70. Lophius budegassa Lophiidae Balearic Sea 1976–1978 T M 337 F, W Macpherson (1979a, 1981)
71. Lophius budegassa Lophiidae G. Saronikos 1975–1976 T S 600 F, N, W, ES Tsimenides (1980)
72. Lophius piscatorius Lophiidae Thracian Sea 1975–76 T S 218 F, N, W, ES Tsimenides (1980)
Notacanthiformes
73. Notacanthus bonaparte† Notacanthidae Balearic Sea 1976–1978 T M 181 F, W Macpherson (1979a, 1981)
Ophidiiformes
74. Cataetyx alleni Bythitidae Balearic Sea July 1987 – 1 16 F, N, ES Carrassón and Matallanas (1990)
75. Ophidion barbatum Ophidiidae Catalan Sea – – M 422 F, N, W, ES Matallanas (1980)
76. Ophidion barbatum Ophidiidae G. Marseille 1980 T M 1 F, N, W, ES Bell and Harmelin-Vivien (1983)
77. Parophidion vassali Ophidiidae G. Lions June 1981 T 1 14 F, N, W, ES Khoury (1984)
Osmeriformes
78. Alepocephalus rostratus Alepocephalidae Balearic Sea July 1987 – 1 128 F, N, ES Carrassón and Matallanas (1990)
79. Alepocephalus rostratus Alepocephalidae Balearic Sea 1987–1991 T S 430 F, N, W, ES Carrassón and Matallanas (1998)
Perciformes
80. Apogon imberbis Apogonidae Corsica – – – – W Pinnegar and Polunin (2000)
81. Parablennius gattorugine Blenniidae G. Lions June 1979 SC 1 4 F, N Zander and Berg (1984)
82. Parablennius rouxi Blenniidae G. Lions June 1979 SC 1 11 F, N Zander and Berg (1984)
83. Parablennius tentacularis† Blenniidae G. Marseille 1980 T M 4 F, N, W, ES Bell and Harmelin-Vivien (1983)
84. Callionymus risso Callionymidae G. Marseille 1980 T M 1 F, N, W, ES Bell and Harmelin-Vivien (1983)
85. Seriola dumerili Carangidae Catalan Sea 1989–1991 T S 385 F, N, W, ES Matallanas et al. (1995)
86. Seriola dumerili Carangidae Tyrrhenian Sea 1992 H, GN M 168 F, ES Pipitone and Andaloro (1995)
87. Seriola dumerili Carangidae C Mediterranean 1989–1992 PS 1 308 F, N, W, ES Andaloro and Pipitone (1997)
88. Seriola dumerili Carangidae G. Castellammare 1990 TN, PS M 60 F, N, W, ES Badalamenti et al. (1995)
89. Seriola dumerili Carangidae G. Castellammare 1992 PS M 166 F, N, W, ES Mazzola et al. (1993)
90. Trachurus mediterraneus Carangidae G. Lions 1977 – – 55 F Ben Salem (1988)
91. Trachurus mediterraneus Carangidae Tunisia 1978 – – 86 F Ben Salem (1988)
92. Trachurus mediterraneus Carangidae Aegean Sea 1978 – – 37 F Ben Salem (1988)
93. Trachurus mediterraneus Carangidae C Aegean Sea 1979–1980 – – 985 N, W Kyrtatos (1998a, b)
94. Trachurus trachurus Carangidae G. Lions 1978 – 1 52 F Ben Salem (1988)
95. Trachurus trachurus Carangidae Tunisia 1978 – 1 78 F Ben Salem (1988)
96. Trachurus trachurus Carangidae Aegean Sea 1978 – 1 35 F Ben Salem (1988)
97. Trachurus trachurus Carangidae Cyclades Islands – – – – RA Kyrtatos (1982)
98. Spicara maena Centracanthidae G. Lions June 1981 T 1 46 F, N, W, ES Khoury (1984)
99. Spicara maena Centracanthidae Corsica – – – – W Pinnegar and Polunin (2000)
100. Spicara maena† Centracanthidae G. Patraikos 1984–1985 T S 355 F, N, W, ES Mytilineou (1987)
101. Spicara smaris Centracanthidae G. Marseille 1980 T M 6 F, N, W, ES Bell and Harmelin-Vivien (1983)
102. Spicara smaris Centracanthidae G. S Evvoikos 1992–1993 GN, TN S 14 F, ES Petrakis et al. (1993)
103. Cepola macrophthalma Cepolidae G. Evvoikos 1986–1988 T S 493 N, ES Stergiou (1993)
104. Epigonus telescopus Epigonidae Balearic Sea 1976–1978 T M 311 F, W Macpherson (1981)
105. Buenia jeffreysii† Gobiidae G. Lions June 1979 SC 1 8 F, N Zander and Berg (1984)
106. Deltentosteus quadrimaculatus Gobiidae G. Lions June 1979 SC 1 32 F, N Zander and Berg (1984)
107. Deltentosteus quadrimaculatus Gobiidae G. Marseille 1980 T M 21 F, N, W, ES Bell and Harmelin-Vivien (1983)
108. Deltentosteus quadrimaculatus Gobiidae Cretan waters 1990–1992 T M – W Labropoulou and Markakis (1998)
109. Gobius auratus Gobiidae G. Lions June 1979 SC 1 14 F, N Zander and Heymer (1992)
110. Gobius auratus Gobiidae G. Lions 1987 SC 2 25 F, N Zander and Berg (1984)
111. Gobius bucchichi Gobiidae N Adriatic Sea – – – 157 W Pölzer and Patzner (1998)
112. Gobius cruentatus Gobiidae G. Marseille 1980 T M 29 F, N, W, ES Bell and Harmelin-Vivien (1983)
113. Gobius fallax Gobiidae G. Marseille 1980 T M 6 F, N, W, ES Bell and Harmelin-Vivien (1983)
114. Gobius geniporus Gobiidae G. Lions 1987 SC 2 125 F, N Zander and Heymer (1992)
115. Gobius niger Gobiidae G. Marseille 1980 T M 15 F, N, W, ES Bell and Harmelin-Vivien (1983)
116. Gobius niger Gobiidae C Adriatic Sea 1979–1980 T M 427 F, N, ES Fabi and Froglia (1983a, b)
117. Gobius niger Gobiidae Cretan waters 1990–1992 T M 328 F, N, W, ES Labropoulou and Markakis (1998);
Labropoulou and Papadopoulou-Smith (1999)
222
Table 1A. Continued

Species Family Area Year SM SF N Method Reference

118. Pomatoschistus bathi Gobiidae G. Lions June 1979 SC 1 25 F, N Zander and Berg (1984)
119. Pomatoschistus bathi Gobiidae G. Lions 1987 SC 2 25 F, N Zander and Heymer (1992)
120. Pomatoschistus quagga Gobiidae G. Marseille 1980 T M 1 F, N, W, ES Bell and Harmelin-Vivien (1983)
121. Zosterisessor ophiocephalus Gobiidae G. Marseille 1980 T M 9 F, N, W, ES Bell and Harmelin-Vivien (1983)
122. Coris julis Labridae G. Lions June 1981 T 1 16 F, N, W, ES Khoury (1984)
123. Coris julis Labridae G. Marseille 1980 T M 21 F, N, W, ES Bell and Harmelin-Vivien (1983)
124. Coris julis Labridae Corsica – – – – W Pinnegar and Polunin (2000)
125. Coris julis Labridae G. S Evvoikos 1992–1993 GN, TN S 81 F, ES Petrakis et al. (1993)
126. Ctenolabrus rupestris Labridae G. Marseille 1980 T M 10 F, N, W, ES Bell and Harmelin-Vivien (1983)
127. Labrus merula Labridae G. Marseille 1980 T M 4 F, N, W, ES Bell and Harmelin-Vivien (1983)
128. Labrus viridis Labridae G. Marseille 1980 T M 14 F, N, W, ES Bell and Harmelin-Vivien (1983)
129. Symphodus cinereus Labridae G. Lions June 1981 T 1 34 F, N, W, ES Khoury (1984)
130. Symphodus cinereus Labridae G. Marseille 1980 T M 75 F, N, W, ES Bell and Harmelin-Vivien (1983)
131. Symphodus cinereus Labridae G. S Evvoikos 1992–1993 GN, TN S 48 F, ES Petrakis et al. (1993)
132. Symphodus mediterraneus Labridae G. Marseille 1980 T M 18 F, N, W, ES Bell and Harmelin-Vivien (1983)
133. Symphodus mediterraneus Labridae G. S Evvoikos 1992–1993 GN, TN S 19 F, ES Petrakis et al. (1993)
134. Symphodus melanocercus Labridae G. Marseille 1980 T M 19 F, N, W, ES Bell and Harmelin-Vivien (1983)
135. Symphodus ocellatus Labridae G. Lions June 1981 T 1 122 F, N, W, ES Khoury (1984)
136. Symphodus ocellatus Labridae G. Marseille 1980 T M 31 F, N, W, ES Bell and Harmelin-Vivien (1983)
137. Symphodus ocellatus Labridae Corsica – – – – W Pinnegar and Polunin (2000)
138. Symphodus ocellatus Labridae G. S Evvoikos 1992–1993 GN, TN S 30 F, ES Petrakis et al. (1993)
139. Symphodus rostratus Labridae G. Lions June 1981 T 1 61 F, N, W, ES Khoury (1984)
140. Symphodus rostratus Labridae G. Marseille 1980 T M 68 F, N, W, ES Bell and Harmelin-Vivien (1983)
141. Symphodus rostratus Labridae G. S Evvoikos 1992–1993 GN, TN S 25 F, ES Petrakis et al. (1993)
142. Symphodus tinca Labridae G. Marseille 1980 T M 23 F, N, W, ES Bell and Harmelin-Vivien (1983)
143. Symphodus tinca Labridae Corsica – – – – W Pinnegar and Polunin (2000)
144. Symphodus tinca Labridae G. S Evvoikos 1992–1993 GN, TN S 21 F, ES Petrakis et al. (1993)
145. Mullus barbatus Mullidae Tyrrhenian Sea 1977–1978 – M 442 F, N, ES Focardi et al. (1979, 1980)
146. Mullus barbatus Mullidae G. Castellammare 1990–1991 TN M 84 F, W, ES Badalamenti et al. (1993)
147. Mullus barbatus Mullidae G. Castellammare 1994–1995 TN M 162 F, N, W Lipari et al. (1998)
148. Mullus barbatus Mullidae W Adriatic Sea 1975 T – 474 F, W, ES Froglia (1988)
149. Mullus barbatus Mullidae C Adriatic Sea 1963–1964 T M 282 N, ES Jukić (1972)
150. Mullus barbatus Mullidae G. Saronikos 1975–1976 T S 346 F, N, W, ES Caragitsou and Tsimenides (1982a)
151. Mullus barbatus Mullidae G. Thermaikos 1976–1977 T S 564 F, N, W, ES Caragitsou and Tsimenides (1982a)
152. Mullus barbatus Mullidae Thracian Sea 1976–1977 T S 758 F, W, ES Caragitsou and Tsimenides (1982b)
153. Mullus barbatus Mullidae G. Patraikos 1983–1984 T S 375 F, N, W, ES Papaconstantinou and Caragitsou (1987b)
154. Mullus barbatus Mullidae G. Korinthiakos 1983–1984 T S 294 F, N, W, ES Papaconstantinou and Caragitsou (1987b)
155. Mullus barbatus Mullidae Ionian Sea 1983–1984 T S 375 F, N, W, ES Papaconstantinou and Caragitsou (1987b)
156. Mullus barbatus Mullidae G. Amvrakikos 1986–1987 TN M 403 F, N, W, ES Vassilopoulou and Papaconstantinou (1993a)
157. Mullus barbatus Mullidae C Aegean Sea 1990–1992 T S 587 F, N, ES Vassilopoulou and Papaconstantinou (1993b)
158. Mullus barbatus Mullidae Cretan waters 1990–1992 T M 630 F, N, W, ES Labropoulou and Papadopoulou-Smith (1999)
Labropoulou and Eleftheriou (1997)
159. Mullus barbatus Mullidae G. S Evvoikos 1992–1993 GN, TN S 70 F, ES Petrakis et al. (1993)
160. Mullus barbatus Mullidae SE Mediterranean 1983–1986 T – 203 F, N, W, ES Golani (1994); Golani and Galil (1991)
161. Mullus surmuletus Mullidae G. Marseille 1980 T M 19 F, N, W, ES Bell and Harmelin-Vivien (1983)
162. Mullus surmuletus Mullidae Corsica – – – – W Pinnegar and Polunin (2000)
163. Mullus surmuletus Mullidae G. Palermo 1981–1982 TN M 232 F, N, W, ES Arculeo et al. (1989a)
164. Mullus surmuletus Mullidae G. Palermo 1981–1983 TN M 232 F, N, W, ES Arculeo et al. (1989b)
165. Mullus surmuletus Mullidae G. Castellammare 1990–1991 TN M 84 F, W, ES Badalamenti et al. (1993)
166. Mullus surmuletus Mullidae Cyclades Islands – – – – RA Kyrtatos (1982)
167. Mullus surmuletus Mullidae Cretan waters 1988–1989 T S 429 N, W Labropoulou and Eleftheriou (1997)
168. Mullus surmuletus Mullidae Cretan waters 1990–1992 T M 322 F, N, W, ES Labropoulou et al. (1997)
169. Mullus surmuletus Mullidae Cretan waters 1990–1992 T M 383 F, N, W, ES Labropoulou and Papadopoulou-Smith (1999)
170. Mullus surmuletus Mullidae N Aegean Sea 1992–1993 T S 100 F, N, W, ES Papaconstantinou et al. (1994)
171. Mullus surmuletus Mullidae SE Mediterranean 1983–1986 T – – F, N, W, ES Golani (1994); Golani and Galil (1991)
172. Upeneus asymmetricus Mullidae SE Mediterranean 1983–1986 T – 148 F, N, W, ES Golani and Galil (1991)
173. Upeneus moluccensis Mullidae SE Mediterranean 1983–1986 T – 276 F, N, W, ES Golani (1994); Golani and Galil (1991)
174. Upeneus moluccensis Mullidae SE Mediterranean 1991–1992 T M 711 F, N, ES Kaya et al. (1999)
175. Upeneus pori Mullidae SE Mediterranean 1983–1986 T – 148 F, N, W, ES Golani (1994)
176. Chromis chromis Pomacentridae G. Lions June 1981 T 1 337 F, N, W, ES Khoury (1984)
177. Chromis chromis Pomacentridae G. Marseille 1980 T M 22 F, N, W, ES Bell and Harmelin-Vivien (1983)
178. Sciaena umbra Sciaenidae N Adriatic Sea 1987–1991 N – 349 F, N, W, ES Froglia and Gramitto (1998)
179. Umbrina cirrosa Sciaenidae N Adriatic Sea 1987–1991 N – 536 F, N, W, ES Froglia and Gramitto (1998)
180. Euthynnus alletteratus Scombridae Cyclades Islands – – – – RA Kyrtatos (1982)
181. Sarda sarda Scombridae Cyclades Islands – – – – RA Kyrtatos (1982)
182. Scomber scombrus Scombridae Cyclades Islands – – – – RA Kyrtatos (1982)
183. Thunnus thynnus Scombridae G. Valencia 1989 H M 82 F, N, ES Sanz Brau (1990)
184. Thunnus thynnus Scombridae Ligurian Sea – T – 67 F, N, W, ES Relini Orsi et al. (1995)
185. Thunnus thynnus Scombridae Adriatic Sea 1957–1960 PS S 1176 N, ES Morović (1961)
 223
Table 1A. Continued

Species Family Area Year SM SF N Method Reference

186. Thunnus thynnus Scombridae Cyclades Islands – – – – RA Kyrtatos (1982)

187. Epinephelus aeneus Serranidae Cyclades Islands – – – – RA Kyrtatos (1982)
188. Epinephelus caninus Serranidae Cyclades Islands – – – – RA Kyrtatos (1982)
189. Epinephelus fasciatus† Serranidae Cyclades Islands – – – – RA Kyrtatos (1982)
190. Epinephelus marginatus Serranidae G. Annaba 1993–1994 N M 68 F, N, W, ES Derbal and Kara (1996)
191. Serranus cabrilla Serranidae G. Marseille 1980 T M 11 F, N, W, ES Bell and Harmelin-Vivien (1983)
192. Serranus cabrilla Serranidae Cretan waters 1990–1992 T M 601 N, W Labropoulou and Eleftheriou (1997)
193. Serranus cabrilla Serranidae G. S Evvoikos 1992–1993 GN, TN S 23 F, ES Petrakis et al. (1993)
194. Serranus cabrilla Serranidae Cyclades Islands – – – – RA Kyrtatos (1982)
195. Serranus hepatus Serranidae G. Marseille 1980 T M 4 F, N, W, ES Bell and Harmelin-Vivien (1983)
196. Serranus hepatus Serranidae Cretan waters 1990–1992 T M 583 N, W Labropoulou and Eleftheriou (1997)
197. Serranus hepatus Serranidae Cretan waters 1990–1992 T M 1045 F, N, W, ES Labropoulou et al. (1998)
198. Serranus hepatus Serranidae G. Thermaikos 1993 T S 124 F, N, ES Wagué (1997)
199. Serranus scriba Serranidae S Tyrrhenian Sea 1981–1983 TN M 244 F, N, W, ES Arculeo et al. (1989a, 1993)
200. Serranus scriba Serranidae G. S Evvoikos 1992–1993 GN, TN S 34 F, ES Petrakis et al. (1993)
201. Siganus luridus Siganidae Dodecanese 1985–1986 TN S 209 F, W, ES Stergiou (1988)
202. Siganus luridus Siganidae SE Mediterranean – TN, H – 110 W Lundberg and Golani (1995)
203. Siganus rivulatus Siganidae SE Mediterranean – TN, H – 418 W Lundberg and Golani (1995)
204. Boops boops Sparidae G. Marseille 1980 T M 22 F, N, W, ES Bell and Harmelin-Vivien (1983)
205. Boops boops Sparidae C Adriatic Sea 1963–1964 T M 79 N, ES Jukić (1972)
206. Dentex dentex Sparidae Balearic Islands 1993–1995 TN, H M 210 RA Morales-Nin and Moranta (1997)
207. Dentex dentex Sparidae Cyclades Islands – – – – RA Kyrtatos (1982)
208. Diplodus annularis Sparidae G. Marseille 1980 T M 7 F, N, W, ES Bell and Harmelin-Vivien (1983)
209. Diplodus annularis Sparidae G. Castellammare 1990–1991 TN M 172 F, W, ES Badalamenti et al. (1993)
210. Diplodus annularis Sparidae Cyclades Islands – – – – RA Kyrtatos (1982)
211. Diplodus annularis Sparidae G. Lions – N – 512 F, N, W Rosecchi (1987); Rosecchi and Nouaze (1987)
212. Diplodus puntazzo Sparidae Balearic Sea 1992–1993 Sp. – 16 F, W, ES Sala and Ballesteros (1997)
213. Diplodus puntazzo Sparidae W Sicily 1991 T 1 – F Mirto et al. (1994)
214. Diplodus sargus sargus† Sparidae Balearic Sea 1992–1993 Sp. – 78 F, W, ES Sala and Ballesteros (1997)
215. Diplodus sargus sargus† Sparidae G. Lions – N – 471 F, N, W Rosecchi (1987); Rosecchi and Nouaze (1987)
216. Diplodus sargus sargus† Sparidae G. Castellammare – N M 97 F, N, W Pepe et al. (1998)
217. Diplodus vulgaris Sparidae Balearic Sea 1992–1993 Sp. – 46 F, W, ES Sala and Ballesteros (1997)
218. Diplodus vulgaris Sparidae G. Marseille 1980 T M 25 F, N, W, ES Bell and Harmelin-Vivien (1983)
219. Diplodus vulgaris Sparidae G. Lions – N – 101 F, N, W Rosecchi (1987); Rosecchi and Nouaze (1987)
220. Diplodus vulgaris Sparidae G. Castellammare – H M 128 F, N, W Pepe et al. (1996)
221. Diplodus vulgaris Sparidae Adriatic Sea 1981–1982 – – 103 RA Jardas and Pallaoro (1990)
222. Lithognathus mormyrus Sparidae G. Castellammare 1990–1991 TN M 66 F, W, ES Badalamenti et al. (1993)
223. Lithognathus mormyrus Sparidae Cyclades Islands – – – – RA Kyrtatos (1982)
224. Oblada melanura Sparidae G. Lions 1996 – 1 39 F, N Lenfant and Olive (1998)
225. Pagellus acarne Sparidae G. Evvoikos 1987 T S 400 F, N, W, ES Papaconstantinou et al. (1989)
226. Pagellus bogaraveo Sparidae N Aegean Sea 1992–1993 T S 146 F, N, W, ES Papaconstantinou et al. (1994)
227. Pagellus erythrinus Sparidae Balearic Sea – – – 70 RA Larraneta (1964)
228. Pagellus erythrinus Sparidae G. Lions 1979–1980 N M 230 F, N, W Rosecchi (1983)
229. Pagellus erythrinus Sparidae G. Lions – N – 204 F, N, W Rosecchi and Nouaze (1987)
230. Pagellus erythrinus Sparidae C Tyrrhenian Sea 1981–1982 T M 596 F, W, ES Ardizzone and Messina (1983)
231. Pagellus erythrinus Sparidae C Mediterranean 1982–1983 T 2 310 N, ES Andaloro and Giarritta (1985)
232. Pagellus erythrinus Sparidae C Adriatic Sea – – – 93 RA Rijavec and Županović (1965)
233. Pagellus erythrinus Sparidae C Adriatic Sea 1963–1964 T M 259 N, ES Jukić (1972)
234. Pagellus erythrinus Sparidae G. Saronikos 1978–1979 T S 194 F, N, W, ES Caragitsou and Papaconstantinou (1985)
235. Pagellus erythrinus Sparidae G. Korinthiakos 1983–1984 T S 76 F, N, W, ES Caragitsou and Papaconstantinou (1988)
236. Pagellus erythrinus Sparidae G. Patraikos 1983–1984 T S 140 F, N, W, ES Caragitsou and Papaconstantinou (1988)
237. Pagellus erythrinus Sparidae Ionian Sea 1983–1984 T S 209 F, N, W, ES Caragitsou and Papaconstantinou (1988)
238. Pagellus erythrinus Sparidae Cyclades Islands – – – – RA Kyrtatos (1982)
239. Pagrus auriga Sparidae Tunisia 1982–1983 – – 70 F, N, W, ES Chakroun–Marzouk and Kartas (1987)
240. Pagrus caeruleostictus Sparidae Tunisia 1982–1983 – – 359 F, N, W, ES Chakroun–Marzouk and Kartas (1987)
241. Pagrus pagrus Sparidae Tunisia 1982–1983 – – 150 F, N, W, ES Chakroun-Marzouk and Kartas (1987)
242. Pagrus pagrus Sparidae Dodecanese 1985–1986 H S 122 F, N, W, ES Papaconstantinou and Caragitsou (1989)
243. Pagrus pagrus Sparidae Cretan waters 1988–1991 T S 634 N, W, ES Labropoulou et al. (1999)
244. Pagrus pagrus Sparidae Cretan waters 1990–1992 T M 634 F, N, W, ES Labropoulou and Markakis (1998);
Labropoulou and Papadopoulou-Smith (1999)
245. Pagrus pagrus Sparidae Cyclades Islands – – – – RA Kyrtatos (1982)
246. Sarpa salpa Sparidae G. Lions – N, Sp. – 32 RA Verlaque (1985, 1990)
247. Sarpa salpa Sparidae G. Marseille – N, Sp. – 25 RA Verlaque (1985, 1990)
248. Sarpa salpa Sparidae Corsica – N, Sp. – 75 RA Verlaque (1985, 1990)
249. Sarpa salpa Sparidae W Sicily 1991 T 1 – F Mirto et al. (1994)
250. Sarpa salpa Sparidae S Adriatic Sea 1989 – M – RA Antolić et al. (1984)
251. Sparus auratus† Sparidae G. Lions – N – 183 F, N, W Rosecchi (1987); Rosecchi and Nouaze (1985)
252. Spondyliosoma cantharus Sparidae G. Marseille 1980 T M 1 F, N, W, ES Bell and Harmelin-Vivien (1983)
224
Table 1A. Continued

Species Family Area Year SM SF N Method Reference

253. Lepidopus caudatus Trichiuridae Balearic Sea 1976–1978 T M 145 F, W Macpherson (1979a, c, 1981)
254. Tripterygion delaisi† Tripterygiidae G. Lions June 1979 SC 1 18 F, N Zander and Berg (1984)
255. Tripterygion delaisi† Tripterygiidae G. Lions 1987 SC 2 40 F, N Zander and Heymer (1992)
256. Uranoscopus scaber Uranoscopidae G. Valencia 1982–1983 T M 250 N, ES Sanz (1985)
257. Uranoscopus scaber Uranoscopidae Cyclades Islands – – – – RA Kyrtatos (1982)
258. Xiphias gladius Xiphiidae G. Annaba 1990–199 1 HM 29 F, N, ES Chalabi and Ifrene (1992)
259. Xiphias gladius Xiphiidae Ligurian Sea – T – 129 F, N, W, ES Relini Orsi et al. (1995)
Pleuronectiformes
260. Arnoglossus laterna Bothidae E Mediterranean 1985 T – 296 F, N, ES Avsar (1994)
261. Arnoglossus thori Bothidae G. Marseille 1980 T M 1 F, N, W, ES Bell and Harmelin–Vivien (1983)
262. Bothus podas Bothidae G. Marseille 1980 T M 2 F, N, W, ES Bell and Harmelin-Vivien (1983)
263. Bothus podas Bothidae C Mediterranean 1985–1987 T S 111 F Schintu et al. (1994)
264. Citharus linguatula Citharidae Cretan waters 1990–1992 T M – W Labropoulou and Markakis (1998)
265. Symphurus nigrescens Cynoglossidae Balearic Sea 1979–1977 T M 1256 F, W Macpherson (1978c, 1979a)
266. Symphurus nigrescens Cynoglossidae Balearic Sea 1976–1978 T M 1256 F, W Macpherson (1981)
267. Lepidorhombus boscii Scophthalmidae Balearic Sea 1976–78 T M 669 F, W Macpherson (1979a, c, 1981)
268. Lepidorhombus boscii Scophthalmidae G. Valencia 1991–1994 CL – 344 F, N, W, ES Morte et al. (1999)
269. Lepidorhombus boscii Scophthalmidae Tyrrhenian Sea 1985–1988 T S 478 N Mannini et al. (1990)
270. Lepidorhombus boscii Scophthalmidae Tyrrhenian Sea 1990–1992 T S 347 F, N, W, ES Sartor and De Ranieri (1996)
271. Lepidorhombus boscii Scophthalmidae N Aegean Sea 1990–1992 T S 804 F, N Papaconstantinou et al. (1993)
272. Lepidorhombus whiffiagonis Scophthalmidae G. Valencia 1991–94 CL – 159 F, N, W, ES Morte et al. (1999)
273. Buglossidium luteum† Soleidae Adriatic Sea 1968–1970 T M – – Giovanardi and Piccinetti (1981)
274. Solea solea Soleidae Balearic Sea 1976–1978 – – 365 F, N, ES Ramos (1981)
275. Solea solea Soleidae Balearic Sea 1987 – M 461 F, N, ES Molinero and Flos (1991)
Rajiformes
276. Dasyatis chrysonata marmorata† Dasyatidae Tunisia 1975–1979 – – 401 F, ES Capape and Zaouali (1992)
277. Raja miraleuts Rajidae Egypt – CL – 381 F Ezzat et al. (1987)
278. Raja radula Rajidae Cyclades Islands – – – – RA Kyrtatos (1982)
279. Rhinobatos rhinobatos Rhinobatidae Egypt 1989–1990 T M 233 F, N, W, ES Abdel-Aziz et al. (1993)
Scorpaeniformes
280. Scorpaena notata Scorpaenidae G. Lions June 1981 T 1 37 F, N, W, ES Khoury (1984)
281. Scorpaena notata Scorpaenidae G. Marseille 1980 T M 35 F, N, W, ES Bell and Harmelin-Vivien (1983)
282. Scorpaena notata Scorpaenidae G. Lions 1980–1986 T – 123 F, N, W, ES Harmelin-Vivien et al. (1989)
283. Scorpaena porcus Scorpaenidae G. Lions June 1981 T 1 133 F, N, W, ES Khoury (1984)
284. Scorpaena porcus Scorpaenidae G. Gabès – – S 798 F, N, W, ES Bradai and Bouain (1990)
285. Scorpaena porcus Scorpaenidae G. Lions 1980–1986 T – 139 F, N, W, ES Harmelin-Vivien et al. (1989)
286. Scorpaena porcus Scorpaenidae G. Marseille 1980 T M 30 F, N, W, ES Bell and Harmelin-Vivien (1983)
287. Scorpaena porcus Scorpaenidae G. Palermo 1981–1983 TN M 321 F, N, W, ES Arculeo et al. (1989a, 1993)
288. Scorpaena porcus Scorpaenidae C Adriatic Sea 1987–1991 TN S 298 F, N, W, ES Pallaoro and Jardas (1990)
289. Scorpaena porcus Scorpaenidae G. S Evvoikos 1992–1993 GN, TN S 19 F, ES Petrakis et al. (1993)
290. Scorpaena porcus Scorpaenidae Cyclades Islands – – – – RA Kyrtatos (1982)
291. Scorpaena scrofa Scorpaenidae G. Marseille 1980 T M 2 F, N, W, ES Bell and Harmelin-Vivien (1983)
292. Scorpaena scrofa Scorpaenidae G. Lions 1980–86 T – 12 F, N, W, ES Harmelin-Vivien et al. (1989)
293. Scorpaena scrofa Scorpaenidae G. Gabès – – S 356 F, N, W, ES Bradai and Bouain (1990)
294. Scorpaena scrofa Scorpaenidae Cyclades Islands – – – – RA Kyrtatos (1982)
295. Helicolenus dactylopterus Sebastidae Balearic Sea 1976–1978 T M 808 F, W Macpherson (1979a, c, 1981)
296. Helicolenus dactylopterus Sebastidae Ionian Sea 1996–1997 T M 382 F, N, ES Madurell and Labropoulou (2000)
297. Chelidonichthys cuculus† Triglidae Catalan Sea 1985–1986 T S 193 F, N, W, ES Moreno-Amich (1992)
298. Chelidonichthys cuculus† Triglidae C Tyrrhenian Sea 1985–1987 T S 70 F, N, ES Colloca et al. (1994)
299. Chelidonichthys gurnardus† Triglidae Catalan Sea 1985–1986 T S 589 F, N, W, ES Moreno-Amich (1994)
300. Chelidonichthys lastoviza† Triglidae G. Marseille 1980 T M 1 F, N, W, ES Bell and Harmelin-Vivien (1983)
301. Chelidonichthys lastoviza† Triglidae Cretan waters 1988–1991 T M 396 N, W, ES Labropoulou and Machias (1998)
302. Chelidonichthys lastoviza† Triglidae Cretan waters 1990–1992 T M 346 F, N, ES Labropoulou and Plaitis (1995)
303. Chelidonichthys lucerna† Triglidae G. Valencia 1989–1991 – – 208 F, N, ES Morte et al. (1997)
304. Chelidonichthys lucerna† Triglidae C Tyrrhenian Sea 1985–1987 T S 135 F, N, ES Colloca et al. (1994)
305. Chelidonichthys lucerna† Triglidae W Adriatic Sea 1973–1974 T S 310 F, N Froglia (1976)
306. Chelidonichthys obscurus† Triglidae Catalan Sea 1985–1986 T S 372 F, N, W, ES Moreno-Amich (1996)
307. Chelidonichthys obscurus† Triglidae G. Valencia 1989–1991 – – 231 F, N, ES Morte et al. (1997)
308. Chelidonichthys obscurus† Triglidae C Tyrrhenian Sea 1985–1987 T S 75 F, N, ES Colloca et al. (1994)
309. Lepidotrigla cavillone Triglidae Catalan Sea 1978–1979 GN M 831 F, N, W, ES Moreno and Matallanas (1983)
310. Lepidotrigla cavillone Triglidae C Tyrrhenian Sea 1985–1987 T S 141 F, N, ES Colloca et al. (1994)
311. Lepidotrigla cavillone Triglidae G. Saronikos 1977–1978 T S 437 F, N, W, ES Caragitsou and Papaconstantinou (1990)
312. Lepidotrigla cavillone Triglidae G. Pagassitikos 1977–1978 T S 270 F, N, W, ES Caragitsou and Papaconstantinou (1990)
313. Lepidotrigla cavillone Triglidae G. Thermaikos 1977–1978 T S 303 F, N, W, ES Caragitsou and Papaconstantinou (1990)
314. Lepidotrigla cavillone Triglidae Thracian Sea 1977–1978 T S 427 F, N, W, ES Caragitsou and Papaconstantinou (1990)
315. Lepidotrigla cavillone Triglidae Cretan waters 1988–1991 T S 694 N, W, ES Labropoulou and Machias (1998)
 225
Table 1A. Continued

Species Family Area Year SM SF N Method Reference

316. Trigla lyra Triglidae Balearic Sea 1976–1978 T M 230 F, W Macpherson (1979a, c, 1981)
317. Trigla lyra Triglidae S Adriatic Sea 1961 – – 234 N Jardas and Županović (1983)
318. Trigla lyra Triglidae G. Saronikos 1989 T M 246 F, N, W, ES Caragitsou and Papaconstantinou (1994)
319. Trigla lyra Triglidae N Aegean Sea 1990–1992 T S 185 N, W, ES Papaconstantinou et al. (1993)
Squaliformes
320. Centroscymnus coelolepis Dalatiidae Catalan Sea 1988–1990 T, H – 86 F, N, W, ES Carrassón et al. (1992)
321. Dalatias licha† Dalatiidae Balearic Sea 1976–1977 T M 31 F, W Macpherson (1979a, 1980a, 1981)
322. Etmopterus spinax Dalatiidae Balearic Sea 1976–1977 T M 353 F, W Macpherson (1979a, 1980a)
323. Etmopterus spinax Dalatiidae Balearic Sea 1976–1978 T M 355 F, W Macpherson (1981)
Syngnathiformes
324. Hippocampus hippocampus Syngnathidae G. Marseille 1980 T M 2 F, N, W, ES Bell and Harmelin-Vivien (1983)
325. Syngnathus acus Syngnathidae G. Lions June 1981 T 1 17 F, N, W, ES Khoury (1984)
326. Syngnathus acus Syngnathidae G. Marseille 1980 T M 21 F, N, W, ES Bell and Harmelin-Vivien (1983)
327. Syngnathus typhle Syngnathidae G. Marseille 1980 T M 14 F, N, W, ES Bell and Harmelin-Vivien (1983)
Torpediniformes
328. Torpedo marmorata Torpedinidae Egypt 1991–1992 T M 84 F, N, W, ES Abdel-Aziz (1994)
329. Torpedo torpedo Torpedinidae Egypt 1991–1992 T M 177 F, N, W, ES Abdel–Aziz (1994)
Zeiformes
330. Capros aper Caproidae Balearic Sea 1976–1978 T M 1067 F, W Macpherson (1979a, c, 1981)
331. Zeus faber Zeidae G. Marseille 1980 T M 1 F, N, W, ES Bell and Harmelin-Vivien (1983)
332. Zeus faber Zeidae G. Evvoikos and Pagassitikos 1986–1988 T S 181 F, N, W, ES Stergiou and Fourtouni (1991)

1995, 2000a; Pauly et al., 1995, 1998c; Pauly and
Palomares, 2000). Real consumers do not usually
have TROPHs with integer values and the definition
of TROPH for any consumer species (i) is (Pauly et
al., 1995, 1998c; Pauly and Christensen 2000a; Pauly
and Palomares, 2000):


G j =1
TROPHi = 1 + DCij ∗ TROPHj ,
j =1
where TROPHj is the fractional trophic level of prey
(j), DCij represents the fraction of j in the diet of
i and G is the total number of prey species. Thus
defined, the TROPH of aquatic consumers is a measur-
able entity that can take any value between 2.0, for
herbivorous/detrivorous, and 5.0, for piscivorous/
carnivorous organisms (Pauly et al., 1998c; Pauly and
Palomares, 2000).
TROPH values were calculated from each dataset
using TrophLab (Pauly et al., 2000d), which is a
stand-alone application for estimating TROPHs and
their standard errors (SE) from two different types
of data: (a) from quantitative diet composition data
(i.e., using the weight or volume contribution of each
prey species to the diet); and (b) from qualitative diet
composition data (i.e., using only the list of prey items
known to occur in the diet). The latter is employed
when diet composition is expressed using indices that
are not good indicators of how much a particular
item contributes to the diet of a given species (e.g.,
numerical contribution and frequency of occurrence of
prey) (Pauly et al., 2000d). TrophLab also provides an
estimate of omnivory (i.e., species feeding on more
than one trophic level) by estimating an omnivory
index (OI) defined as:


G
OI = (TROPHj − TROPHi )2 ∗ DCij .
The OI value equals zero when feeding on prey of the
same TROPH and increases with an increase in the
variety of prey’s TROPH. The square root of OI is the
SE of the TROPH (Christensen and Pauly, 1992a, b).
TROPHs and their SE (Table 1B) were estimated
from the gravimetric abundances of prey (which were
available for 222 out of a total of 332 datasets)
using the “quantitative approach” of TrophLab. For all
remaining datasets, TROPHs were estimated from the
list of prey items known to occur in the diet using the
“qualitative approach” of TrophLab.
The histogram of all estimated TROPH values was
decomposed into separate normal distributions and the
corresponding mean and standard deviation (SD) of
TROPH per normal distribution were calculated. The
decomposition was done using Bhattacharya’s (1967)
method, which is incorporated into FiSAT (Gayanilo
and Pauly, 1997).
TROPH values from each dataset were regressed
against the Lmax of the species in concern. Finally,
for studies reporting diet composition per length class,
TROPH was also estimated separately for each length
class and its relationship with the mid-point of the
226
Table 1B. Feeding habits of Mediterranean marine fish. Length range: length range (or mean length) of specimens (in cm); W (or N):
gravimetric (or numerical) contribution (%) of prey in the diet of the fish species examined; TROPH: trophic level; SE: standard error of
TROPH; and Lmax : maximum body length (in cm). Habitat type and Lmax extracted from “FishBase online” (www.fishbase.org). ∗∗Diet
composition data expressed as frequency of occurrence (F); † Valid names of species (from “FishBase online”; www.fishbase.org)

Species Length range Main prey W (or N) Habitat TROPH SE Lmax

1. Conger conger – Fish (Micromesistius poutassou, Gadiculus argenteus 51, 16, 33 Bathydemersal 3.20 0.40 300
argenteus), crustaceans, other
2. Conger conger 9–22 Decapods, brachyurans, polychaetes, other 45, 22, 15, 18 Bathydemersal 4.49 0.80 300
3. Conger conger – Fish, polychaetes, decapods 99.5, 0.4, 0.1 Bathydemersal 3.42 0.50 300
4. Gnathophis mystax 10–39 10–19 cm, decapods (Alpheus glaber, Calocaris 90, 4, 6 Demersal 3.55 0.57 60
macandreae), polychaetes, other
20–29 cm, decapods (Alpheus glaber, Calocaris 66, 21, 13 3.85 0.61
macandreae), fish, other
30–39 cm, decapods (Goneplax rhomboides, Processa 92, 5, 3 3.60 0.59
mediterranea), polychaetes, other
5. Gnathophis mystax 13–40 Decapods (Alpheus glaber, Processa canaliculata), fish, 71, 8, 4, 17 Demersal 3.62 0.57 60
mysids, other
6. Ophichthus rufus – Fish, molluscs, decapods, other 58, 16, 16, 10 Demersal 4.25 0.68 60
7. Atherina boyeri Mean TL 3.7 Copepods, ostracods, polychaetes, amphipods, other ∗∗ Demersal 3.30 0.43 20
8. Bathypterois mediterraneus 5–18 Copepods, crustaceans, mysids, decapods, other (32, 23, 19, 11, 15) Bathydemersal 3.20 0.43 19
9. Saurida undosquamis – Fish (Mullus barbatus), cephalopods (99, 1) Demersal 4.49 0.79 50
10. Saurida undosquamis 14–33 Fish (Engraulis encrasicolus), crustaceans, molluscs (95, 4, 1) Demersal 3.80 0.56 50
11. Synodus saurus 16–26 Fish (Boops boops, Solea solea), molluscs, crustaceans (93, 5, 2) Demersal 4.20 0.58 40
12. Sargocentron rubrum – Decapods, fish, polychaetes, molluscs, isopods 81, 6, 6, 4, 3 Reef-associated 3.50 0.53 32
13. Hoplostethus mediterraneus 9–21 Decapods, euphausids, fish, mysids, other (42, 35, 10, 6, 7) Benthopelagic 3.50 0.53 42
14. Galeus melastomus 10–60 Fish (Engraulis encrasicolus, Gadiculus argenteus 52, 23, 19, 6 Bathydemersal 4.26 0.67 61
argenteus), decapods, cephalopods, other
15. Galeus melastomus – Fish (Engraulis encrasicolus, Gadiculus argenteus 42, 12, 11, 35 Bathydemersal 3.70 0.57 61
argenteus), crustaceans, molluscs, other
16. Galeus melastomus 15–61 371–667 m: 15–30 cm, decapods (Sergia robusta, 65, 14, 3, 18 Bathydemersal 3.86 0.65 61
Calocaris macandreae), fish, euphausids, other
40–60 cm, decapods (Pasiphaea multidentate, Calocaris 56, 17, 13, 14 3.80 0.57
macandreae), cephalopods, fish, other
984–1584 m: 10–39 cm, fish (Antonogadus 58, 35, 5, 2 4.17 0.71
megalokynodon), decapods, cephalopods, other
40–61 cm, cephalopods (Todarodes sagittatus), fish, 37, 33, 26, 4 4.18 0.59
decapods, other
17. Scyliorhinus canicula – Fish (Micromesistius poutassou, Gadiculus argenteus 41, 26, 7, 26 Demersal 3.80 0.59 100
argenteus), decapods, molluscs, other
18. Chimaera monstrosa 19–60 19–29 cm, ophiurids, amphipods, polychaetes, other 93, 3, 2, 2 Bathydemersal 3.38 0.35 120
30–39 cm, ophiurids, decapods, polychaetes, other 91, 4, 3, 2 3.39 0.36
40–49 cm, ophiurids, decapods, other 84, 13, 3 3.42 0.39
50–59 cm, ophiurids, decapods, other 59, 35, 6 3.51 0.47
>60 cm, decapods (Medaeus couchi, Polycheles typhlops), 94, 5, 1 3.59 0.58
ophiurids, other
19. Sardina pilchardus 12–16 Copepods, plants, cladocerans ∗∗ Pelagic 3.20 0.42 25
20. Sardina pilchardus 10–14 Diatoms, copepods, euphausids, eggs, larvae, algae (85, 6, 2, 3, 3, 1) Pelagic 3.10 0.32 25
21. Engraulis encrasicolus Mean TL 11.0 Copepods (Centropages typicus, Onceae spp.), molluscs, ∗∗ Pelagic 3.50 0.48 20
other
22. Engraulis encrasicolus Mean TL 12.3 Copepods, cladocerans, crustaceans, appendicularians, ∗∗ Pelagic 3.40 0.44 20
molluscs, other
23. Engraulis encrasicolus Mean TL 13.4 Copepods, cladocerans, crustaceans, appendicularians, ∗∗ Pelagic 3.40 0.44 20
molluscs, other
24. Gadiculus argenteus argenteus – Euphausids, fish, decapods, other 57, 22, 17, 4 Pelagic 3.90 0.64 15
25. Gadiculus argenteus argenteus – Crustaceans (Meganyctiphanes norvegica, Nyctiphanes 74, 7, 19 Pelagic 3.55 0.54 15
couchii), fish, other
26. Micromesistius poutassou 10–35 10–16 cm, fish, euphausids, decapods, other 54, 21, 20, 5 Benthopelagic 3.97 0.64 50
17–23 cm, fish, euphausids, decapods, other 51, 29, 20, 5 3.95 0.67
24–35 cm, fish, decapods, euphausids, cephalopods, other 42, 25, 18, 12 4.04 0.64
27. Micromesistius poutassou – Crustaceans (Meganyctiphanes norvegica, Pasiphaea 39, 26, 35 Benthopelagic 3.90 0.64 50
sivado), fish, other
28. Micromesistius poutassou Mean TL 17.2 <18.5 cm, euphausids, crustaceans, fish, mysids, 76, 11, 5, 4, 3, 1 Benthopelagic 3.34 0.44 50
cephalopods, decapods
>18.5 cm, euphausids, fish, crustaceans, decapods, 45, 38, 5, 4, 4, 4 3.78 0.59
cephalopods, mysids
29. Micromesistius poutassou Mean TL 24.9 Fish, decapods, mysids, isopods 87, 11, 1, 1 Benthopelagic 4.39 0.77 50
30. Micromesistius poutassou 5–27 Euphausids, decapods, fish, cephalopods, other 32, 31, 29, 3, 5 Benthopelagic 3.77 0.60 50
31. Trisopterus minutus† 8–21 Crustaceans (Alpheus glaber, Processa nouveli), fish, 80, 18, 2 Benthopelagic 3.76 0.63 40
other
 227
Table 1B. Continued

Species Length range Main prey W (or N) Habitat TROPH SE Lmax

32. Trisopterus minutus† 4–21 Mysids (Processa nouveli, Alpheus glaber), crustaceans, 63, 11, 9, 17 Benthopelagic 3.39 0.49 40
fish, other
33. Trisopterus minutus† 5–27 Decapods (Alpheus glaber), fish, other 52, 30, 18 Benthopelagic 3.81 0.63 40
34. Trisopterus minutus† 9–23 Fish, decapods, brachyurans, other 34, 27, 12, 26 Benthopelagic 3.79 0.64 40
35. Trisopterus minutus† 5–23 Mysids (Lophogaster typicus), decapods, euphausids, (41, 26, 15, 18) Benthopelagic 3.50 0.48 40
other
36. Gaidropsarus biscayensis† 4–14 Decapods, fish, euphausids, other 68, 19, 6, 7 Benthopelagic 3.72 0.62 40
37. Gaidropsarus biscayensis† 5–21 5–12 cm, decapods (Processa nouveli, Alpheus glaber), 73, 9, 15 Benthopelagic 3.61 0.51 40
fish, other
13–21 cm, decapods (Processa nouveli, Alpheus glaber), 80, 18, 2 3.75 0.61
fish, other
38. Gaidropsarus mediterraneus – Decapods, brachyurans, polychaetes, other 45, 22, 15, 18 Demersal 3.41 0.47 5
39. Gaidropsarus mediterraneus – Decapods, amphipods, brachyurans, other 46, 36, 14, 4 Demersal 3.42 0.50 5
40. Gaidropsarus vulgaris – Decapods, brachyurans, polychaetes, other 45, 22, 15, 18 Demersal 3.49 0.38 6
41. Gaidropsarus vulgaris – Decapods, brachyurans, other 94, 4, 2 Demersal 3.42 0.50 6
42. Molva macrophthalma† – Fish (Gadiculus argenteus argenteus, Antonogadus 100 Demersal 4.50 0.80 90
megalokynodon)
43. Caelorinchus caelorhincus† 5–39 5–9 cm, polychaetes, amphipods, copepods, decapods, 43, 18, 17, 11, 11 Benthopelagic 3.17 0.38 40
other
10–19 cm, polychaetes, decapods, other 66, 23, 11 3.20 0.38
20–29 cm, decapods (Solenocera membranacea), 61, 34, 3, 2 3.45 0.51
polychaetes, fish, other
30–39 cm, decapods (Alpheus glaber), other 99, 1 3.60 0.59
44. Caelorinchus caelorhincus† – Polychaetes, decapods, other 52, 31, 17 Benthopelagic 3.20 0.41 40
45. Hymenocephalus italicus 9–19 Decapods (Solenocera membranacea), mysids, 48, 24, 11, 9, 8 Benthopelagic 3.40 0.51 25
amphipods, copepods, other
46. Hymenocephalus italicus – Euphausids (Processa mediterranea, Solenocera 45, 31, 9, 15 Benthopelagic 3.20 0.38 25
membranacea), decapods, copepods, other
47. Nezumia aequalis 10–39 10–19 cm, polychaetes (Nephthys sp.), decapods, mysids, 33, 30, 14, 8, 15 Benthopelagic 3.29 0.45 36
isopods, other
20–29 cm, decapods (Calocaris macandreae), 35, 32, 14, 5, 14 3.49 0.53
polychaetes, ophiurids, amphipods, other
30–39 cm, polychaetes, isopods, mysids 85, 10, 5 3.09 0.31
48. Nezumia aequalis – Decapods, polychaetes, ophiurids, other 46, 33, 10, 11 Benthopelagic 3.20 0.39 36
49. Trachyrincus scabrus† 10–39 10–19 cm, decapods, other 96, 4 Benthopelagic 3.58 0.58 60
20–29 cm, decapods, polychaetes, other 92, 6, 2 3.56 0.57
30–39 cm, decapods (Alpheus glaber), fish 75, 25 3.83 0.65
50. Trachyrincus scabrus† – Decapods, other 88, 12 Benthopelagic 3.20 0.40 60
51. Trachyrincus scabrus† 6–18 Amphipods, ostracods, polychaetes, decapods, other (37, 29, 13, 13, 8) Benthopelagic 3.50 0.56 60
52. Merluccius merluccius – Fish (Micromesistius poutassou, Lesueuriogobius friesii), 68, 14, 18 Demersal 3.90 0.64 140
crustaceans, other
53. Merluccius merluccius 8–65 8–14 cm, fish (Sardina pilchardus, Cepola rubescens), 55, 17, 15, 13 Demersal 3.98 0.67 140
euphausids, decapods, mysids
14–25 cm, fish (Sardina pilchardus, Cepola rubescens), 78, 12, 6, 4 4.26 0.75
decapods, euphausids, mysids
25–40 cm, fish (Sardina pilchardus, Cepola rubescens), 90, 10 4.41 0.78
decapods
40–65 cm, fish (Sardina pilchardus, Cepola rubescens), 92, 8 4.43 0.79
decapods
54. Merluccius merluccius 4–74 4–8 cm, euphausids, amphipods, cephalopods, decapods, 60, 22, 14, 2, 2 Demersal 3.43 0.44 140
fish
8–13 cm, euphausids, decapods, fish, cephalopods, 79, 9, 6, 4, 2 3.37 0.46
amphipods
13–18 cm, decapods, euphausids, fish 66, 18, 16 3.67 0.60
18–25 cm, decapods, fish 83, 17 3.75 0.63
25–31 cm, fish, decapods 63, 37 4.17 0.73
31–37 cm, fish, decapods 84, 16 4.36 0.77
>37 cm, fish, decapods, cephalopods 73, 22, 5 4.20 0.74
55. Merluccius merluccius – Fish (Engraulis encrasicolus, Sardina pilchardus), (56, 6, 1) Demersal 4.30 0.61 140
decapods, cephalopods, other
56. Merluccius merluccius 8–35 Fish (Engraulis encrasicolus, Sprattus sprattus), (64, 21, 6, 5, 4) Demersal 4.00 0.67 140
decapods, amphipods, mysids, other
57. Merluccius merluccius 15–52 Fish (Trachurus spp., Boops boops), cephalopods, ∗∗ Demersal 4.40 0.65 140
crustaceans
58. Merluccius merluccius – Fish, cephalopods, decapods, mysids, crustaceans 69, 14, 9, 5, 3 Demersal 4.32 0.71 140
59. Merluccius merluccius 4–38 Fish, euphausids, other 96, 3, 1 Demersal 4.45 0.79 140
228
Table 1B. Continued

Species Length range Main prey W (or N) Habitat TROPH SE Lmax

60. Merluccius merluccius 4–38 Fish, cephalopods, euphausids, other 81, 8, 5, 6 Demersal 4.44 0.78 140
61. Merluccius merluccius 4–38 Fish, euphausids, mysids, other 74, 14, 5, 7 Demersal 4.19 0.72 140
62. Merluccius merluccius – Fish (Micromesistius poutassou, Engraulis encrasicolus), – Demersal 4.10 0.60 140
euphausids, decapods, cephalopods
63. Merluccius merluccius 5–19 Fish (Vinciguerria sp., Argyropelecus hemigymnus), 74, 9, 7, 10 Demersal 4.19 0.72 140
mysids, euphausids, other
64. Merluccius merluccius – Decapods, fish, euphausids, other 37, 27, 20, 16 Demersal 3.75 0.61 140
65. Lepidion lepidion 7–23 Crustaceans, cephalopods, polychaetes (96, 2, 2) Bathypelagic 3.30 0.39 34
66. Lepidion lepidion – Crustaceans (Acanthephyra eximia, Calocaris 65, 17, 10, 8 Bathypelagic 3.67 0.59 34
macandreae), fish, foraminiferans, other
67. Phycis blennoides 6–35 Decapods (Alpheus glaber, Calocaris macandreae), fish 77, 23 Benthopelagic 3.77 0.63 11
68. Phycis phycis 15–49 Fish (Boops boops, Chromis chromis), decapods, 66, 21, 8, 5 Benthopelagic 4.09 0.72 65
brachyurans, other
69. Apletodon dentatus – Amphipods, copepods, foraminiferans 56, 37, 7 Demersal 3.19 0.41 4
70. Lophius budegassa – Fish (Gadiculus argenteus argenteus, Phycis blennoides), 65, 19, 16 Bathydemersal 3.90 0.62 100
crustaceans, other
71. Lophius budegassa – Fish, cephalopods, crustaceans 85, 10, 2 Bathydemersal 4.40 0.64 100
72. Lophius piscatorius – Fish, cephalopods, crustaceans 80, 9, 5 Bathydemersal 4.30 0.65 200
73. Notacanthus bonaparte† – Ophiurids, other 94, 6 Bathypelagic 3.40 0.35 –
74. Cataetyx alleni 6–10 Crustaceans, polychaetes, other (73, 25, 2) Bathypelagic 3.10 0.38 12
75. Ophidion barbatum 9–26 Decapods (Processa canaliculata, Alpheus glaber), 61, 27, 9, 3 Demersal 3.56 0.57 25
mysids, fish, other
76. Ophidion barbatum – Decapods, brachyurans, amphipods 61, 23, 16 Demersal 3.47 0.59 25
77. Parophidion vassali – Decapods, brachyurans, polychaetes, other 53, 22, 15, 10 Demersal 3.43 0.43 25
78. Alepocephalus rostratus 11–37 Coelenterata, crustaceans, fish, other (37, 33, 7, 23) Bathydemersal 3.40 0.40 50
79. Alepocephalus rostratus – 1000–1425 m: Decapods (Sergia robusta, Aristeus 50, 38, 4, 8 Bathydemersal 3.52 0.53
antennatus), pyrosomida, euphausids, others
1425–2250 m: <22 cm, pyrosomida, (Pyrosoma 89, 5, 3, 3 3.39 0.46
atlanticum), gastropods, mysids, other
>22 cm, fish, decapods, cephalopods, pyrosomida, 24, 24, 20, 15, 3, 14 3.91 0.58
gastropods, other
80. Apogon imberbis – Decapods, fish 58, 42 Demersal 3.98 0.69 15
81. Parablennius gattorugine Mean TL 13.3 Algae, sponges, decapods, ascidians, polychaetes, other (40, 30, 14, 6, 3, 7) Demersal 3.00 0.32 30
82. Parablennius rouxi Mean TL 5.7 Algae, balanoids, polychaetes, copepods, decapods, other (40, 28, 10, 8, 6, 8) Demersal 3.20 0.34 8
83. Parablennius tentacularis† – Polychaetes, bivalves, echinoderms, other 54, 35, 5, 6 Demersal 3.11 0.29 15
84. Callionymus risso – Polychaetes, bivalves, foraminiferans 75, 19, 6 Demersal 3.09 0.28 11
85. Seriola dumerili Mean SL 29.6 Juveniles: Fish (Engraulis encrasicolus, Sardina pilchardus), 98, 1, 1 Reff-associated 4.49 0.80 190
cephalopods, other
Adults: Fish (Sardina pilchardus, Merluccius merluccius), 89, 10, 1 4.49 0.77
cephalopods, other
86. Seriola dumerili Mean SL 91.8 9–19 cm, decapod larvae, copepods, other ∗∗ Reef-associated 3.50 0.50 190
20–24 cm, fish, crustaceans, other ∗∗ 3.70 0.55
25–33 cm, fish, other ∗∗ 4.30 0.71
87. Seriola dumerili 9–33 Fish (Boops boops, Sardina pilchardus), cephalopods 71, 29 Reef-associated 4.50 0.70 190
88. Seriola dumerili Mean SL 109.1 <8 cm, crustaceans, fish, other 74, 18, 8 Reef-associated 3.40 0.43 190
8–12 cm, fish, crustaceans, polychaetes, other 55, 24, 8, 13 3.83 0.66
>12 cm, fish, cephalopods, other 74, 17, 9 4.37 0.72
89. Seriola dumerili 2–30 Fish, crustaceans, other 92, 4, 4 Reef-associated 4.37 0.76 190
90. Trachurus mediterraneus 13–24 Crustaceans, fish, molluscs ∗∗ Pelagic 3.70 0.47 60
91. Trachurus mediterraneus 15–32 Fish, crustaceans, molluscs, algae, polychaetes ∗∗ Pelagic 3.50 0.46 60
92. Trachurus mediterraneus 20–38 Crustaceans, polychaetes, fish ∗∗ Pelagic 3.20 0.44 60
93. Trachurus mediterraneus 8–38 Fish larvae, copepods, mysids, other 39, 20, 11, 30 Pelagic 3.60 0.56 60
94. Trachurus trachurus 16–28 Crustaceans, fish ∗∗ Pelagic 3.70 0.48 70
95. Trachurus trachurus 18–27 Crustaceans, fish, molluscs, algae, polychaetes ∗∗ Pelagic 3.40 0.43 70
96. Trachurus trachurus 20–33 Crustaceans, fish, molluscs ∗∗ Pelagic 3.60 0.55 70
97. Trachurus trachurus – Fish (Atherina boyeri, Boops boops), cephalopods, – Pelagic 3.90 0.57 70
crustaceans, fish larvae
98. Spicara maena – Fish, appendicularians, copepods, cladocerans 76, 16, 6, 2 Demersal 3.16 0.27 25
99. Spicara maena – Copepods, cladocerans 100 Demersal 3.00 0.00 25
100. Spicara maena† – Copepods, appendicularians, other (61, 28, 11) Demersal 3.30 0.39 25
101. Spicara smaris – Copepods, mysids 99, 1 Demersal 3.00 0.04 20
102. Spicara smaris 11–19 Copepods, cladocerans, decapod larvae ∗∗ Demersal 3.10 0.35 20
103. Cepola macrophthalma 11–51 Copepods (Calanus spp., Eucalanus sp.), euphausids, (60, 31, 9) Demersal 3.11 0.27 70
other
104. Epigonus telescopus – Decapods (Meganyctiphanes norvegica, Boreomysis 41, 23, 19, 7, 10 Bathydemersal 3.40 0.42 75
megalops), copepods, fish, tunicates, other
 229

Table 1B. Continued

Species Length range Main prey W (or N) Habitat TROPH SE Lmax

105. Buenia jeffreysii† Mean TL 3.9 Copepods, brachyurans, fish, other (56, 24, 8, 12) Reef-associated 3.60 0.52 6
106. Deltentosteus quadrimaculatus Mean TL 2.8 Copepods, decapods, amphipods, polychaetes, other (62, 12, 10, 10, 6) Demersal 3.30 0.35 8
107. Deltentosteus quadrimaculatus – Polychaetes, amphipods, copepods, other 60, 20, 15, 5 Demersal 3.11 0.32 8
108. Deltentosteus quadrimaculatus – Polychaetes, decapods – Demersal 3.30 0.44 8
109. Gobius auratus Mean TL 5.8 Copepods, decapods, isopods, polychaetes, other (43, 15, 12, 10, 20) Demersal 3.10 0.34 10
110. Gobius auratus 5–9 Copepods, decapods, polychaetes, gastropods, algae, other (64, 12, 5, 5, 1, 13) Demersal 3.30 0.40 10
111. Gobius bucchichi 3–12 Crustaceans, algae, polychaetes, molluscs, other 45, 27, 8, 7, 13 Demersal 2.74 0.33 10
112. Gobius cruentatus – Polychaetes, ophiurids, decapods, crustaceans, other 44, 18, 10, 8, 20 Demersal 3.25 0.36 18
113. Gobius fallax – Amphipods, decapods, mysids, crustaceans 68, 26, 4, 2 Demersal 3.37 0.54 9
114. Gobius geniporus 1–6 Copepods, crustaceans, decapods, other (69, 9, 6, 16) Demersal 3.50 0.47 16
115. Gobius niger – Polychaetes, amphipods, mysids, decapods, other 46, 38, 8, 5, 3 Demersal 3.20 0.41 15
116. Gobius niger 5–17 Polychaetes, amphipods, decapods, other ∗∗ Demersal 3.30 0.46 15
117. Gobius niger 5–14 Decapods, polychaetes (Glycera rouxii, Terebellides 54, 29, 17 Demersal 3.47 0.51 15
stroemi), other
118. Pomatoschistus bathi Mean TL 3.1 Copepods, decapods, other (83, 10, 7) Demersal 3.30 0.41 –
119. Pomatoschistus bathi 1–3 Copepods, other (94, 6) Demersal 3.20 0.22 –
120. Pomatoschistus quagga – Amphipods 100 Demersal 3.29 0.53 –
121. Zosterisessor ophiocephalus – Polychaetes, amphipods, other 54, 41, 5 Demersal 3.16 0.40 25
122. Coris julis – Copepods, gastropods, decapods, crustaceans, other 43, 20, 9, 28 Demersal 3.27 0.41 30
123. Coris julis – Bivalves, gastropods, decapods, crustaceans, other 43, 24, 14, 9, 10 Demersal 3.30 0.42 30
124. Coris julis – Molluscs, crustaceans, cephalopods, decapods, isopods, 49, 16, 11, 9, 6, 5, 4 Demersal 3.63 0.47 30
polychaetes, algae
125. Coris julis 5–17 Molluscs, gastropods, decapods ∗∗ Demersal 3.30 0.41 30
126. Ctenolabrus rupestris – Bivalves, ophiurids, amphipods, decapods, other 69, 16, 9, 4, 2 Demersal 3.19 0.35 18
127. Labrus merula – Brachyurans, decapods, echinoderms, other 66, 18, 12, 4 Demersal 3.47 0.55 45
128. Labrus viridis – Fish, brachyurans, echinoderms, decapods, isopods, 41, 22, 9, 9, 7, 7, 5 Demersal 3.84 0.64 47
polychaetes, other
129. Symphodus cinereus 5–8 Copepods, gastropods, decapods, crustaceans, other 43, 20, 9, 28 Demersal 3.28 0.47 16
130. Symphodus cinereus 2–11 Bivalves, gastropods, amphipods, decapods, other 33, 27, 16, 9, 15 Demersal 3.30 0.42 16
131. Symphodus cinereus 5–9 Crustaceans, euphausids, pycnogonids ∗∗ Demersal 3.10 0.39 16
132. Symphodus mediterraneus – Bivalves, amphipods, polychaetes, other 57, 23, 7, 13 Demersal 3.22 0.41 18
133. Symphodus mediterraneus 6–14 Molluscs, algae, copepods, crustaceans ∗∗ Demersal 3.10 0.40 18
134. Symphodus melanocercus – Bivalves, copepods, amphipods, other 40, 27, 19, 14 Demersal 3.25 0.40 14
135. Symphodus ocellatus 6–9 Copepods, crustaceans, echinoderms, gastropods, other 27, 22, 13, 12, 26 Demersal 3.26 0.49 12
136. Symphodus ocellatus – Amphipods, gastropods, copepods, other 38, 36, 20, 6 Demersal 3.30 0.42 12
137. Symphodus ocellatus – Copepods, algae, crustaceans, other 86, 6, 4, 4 Demersal 2.99 0.17 12
138. Symphodus ocellatus 5–10 Crustaceans, pycnogonids ∗∗ Demersal 3.40 0.51 12
139. Symphodus rostratus – Copepods, crustaceans, echinoderms, gastropods, other 27, 22, 13, 12, 26 Demersal 3.41 0.51 13
140. Symphodus rostratus 2–13 Amphipods, decapods, isopods, fish, other 44, 25, 18, 6, 7 Demersal 3.30 0.42 13
141. Symphodus rostratus 5–12 Crustaceans, decapods, mysids ∗∗ Demersal 3.40 0.51 13
142. Symphodus tinca – Amphipods, bivalves, isopods, polychaetes, other 37, 31, 8, 7, 17 Demersal 3.26 0.45 44
143. Symphodus tinca – Polychaetes, crustaceans, algae, decapods, amphipods, 41, 26, 11, 7, 7, 4, 4 Demersal 3.14 0.35 44
molluscs, other
144. Symphodus tinca 13–21 Crustaceans, algae, decapods, molluscs ∗∗ Demersal 3.20 0.44 44
145. Mullus barbatus 9–15 Crustaceans (Ampelisca sp., Leucothea sp.), polychaetes, ∗∗ Demersal 3.30 0.43 30
amphipods, decapods, other
146. Mullus barbatus Mean TL 1.7 Polychaetes, bivalves, algae, amphipods, other ∗∗ Demersal 3.10 0.38 30
147. Mullus barbatus Mean TL 1.7 Polychaetes (Aricidea spp., Megallona rosea), bivalves, 59, 20, 21 Demersal 3.40 0.46 30
other
148. Mullus barbatus 5–9 Decapods (Processa sp.), bivalves, amphipods, 20, 17, 15, 11, 37 Demersal 3.40 0.43 30
polychaetes, other
149. Mullus barbatus – Polychaetes (Sternaspis scutate, Nephthys hystricis), 41, 19, 14, 26 Demersal 3.30 0.47 30
bivalves, crustaceans, other
150. Mullus barbatus 9–19 Decapods (Processa canaliculata, Amolopenaeus 33, 22, 21, 18, 6 Demersal 2.95 0.40 30
elegans), polychaetes, mysids, amphipods, other
151. Mullus barbatus 8–20 Decapods (Crangon crangon, Processa canaliculata), 53, 18, 11 18 Demersal 2.99 0.42 30
polychaetes, mysids, other
152. Mullus barbatus – Decapods (Processa canaliculata, Crangon crangon), 32, 24, 17, 27 Demersal 2.79 0.36 30
polychaetes, mysids, other
153. Mullus barbatus 12–18 Euphausids, polychaetes, mysids, other 34, 34, 20, 12 Demersal 3.48 0.53 30
154. Mullus barbatus 12–18 Polychaetes, euphausids, mysids, other 64, 21, 9, 6 Demersal 3.20 0.38 30
155. Mullus barbatus 12–18 Euphausids, polychaetes, mysids, other 52, 20, 11, 17 Demersal 3.42 0.51 30
156. Mullus barbatus 10–16 Polychaetes (Pectinaria koreni, Sternaspis scutate), 54, 33, 8, 5 Demersal 3.14 0.33 30
molluscs, crustaceans, other
157. Mullus barbatus – Decapods, gastropods, crustaceans (18, 17, 11) Demersal 3.50 0.58 30
230
Table 1B. Continued

Species Length range Main prey W (or N) Habitat TROPH SE Lmax

158. Mullus barbatus 4–17 Polychaetes (Micronepthys maryae, Nepthys assimilis), 52, 21, 21, 6 Demersal 3.31 0.42 30
molluscs, decapods, other
159. Mullus barbatus 9–18 Decapods, amphipods, polychaetes, copepods ∗∗ Demersal 3.50 0.53 30
160. Mullus barbatus – Decapods (Leptochela pugnax, Liocarcinus pusillus), 73, 13, 4, 3, 7 Demersal 3.57 0.57 30
polychaetes, brachyurans, fish, other
161. Mullus surmuletus – Amphipods, mysids, decapods, crustaceans, other 47, 39, 5, 5, 4 Demersal 3.28 0.47 40
162. Mullus surmuletus – Polychaetes, amphipods, decapods, echinoderms, 29, 18, 16, 14, 8, 6, 9 Demersal 3.29 0.43 40
molluscs, isopods, other
163. Mullus surmuletus 10–22 Polychaetes, fish, brachyurans, decapods, other 48, 14, 11, 9, 18 Demersal 3.44 0.49 40
164. Mullus surmuletus 10–22 Polychaetes, fish, brachyurans, decapods, other 49, 16, 10, 9, 16 Demersal 3.38 0.49 40
165. Mullus surmuletus Mean TL 1.8 Polychaetes, bivalves, amphipods, algae, other ∗∗ Demersal 3.20 0.42 40
166. Mullus surmuletus – Decapods, amphipods, brachyurans, bivalves, polychaetes – Demersal 3.30 0.53 40
167. Mullus surmuletus 10–15 Decapods (Inachus sp., Ethusa mascarone), amphipods, 48, 16, 10, 9, 17 Demersal 3.53 0.56 40
fish, polychaetes, other
168. Mullus surmuletus 6–23 Decapods, polychaetes, fish, other 89, 5, 2, 4 Demersal 3.53 0.56 40
169. Mullus surmuletus 6–21 Crustaceans, polychaetes (Aponuphis bilineata, 70, 28, 2 Demersal 3.58 0.58 40
Harmothoe sp.), other
170. Mullus surmuletus 9–12 Decapods (Upogebia tipica, Processa nouveli), 60, 20, 9, 11 Demersal 3.16 0.36 40
brachyurans, polychaetes, other
171. Mullus surmuletus – Decapods (Leptochela pugnax), polychaetes, brachyurans, 58, 29, 5, 2 Demersal 3.51 0.54 40
other
172. Upeneus asymmetricus – Decapods (Leptochela pugnax), crustaceans – Demersal 3.60 0.60 30
173. Upeneus moluccensis – Decapods (Leptochela pugnax, Parapenaeus 41, 38, 11, 4, 6 Demersal 3.40 0.48 20
longirostris), fish, polychaetes, molluscs, other
174. Upeneus moluccensis 8–16 Decapods, copepods, mysids, other (52, 17, 14, 17) Demersal 3.89 0.65 20
175. Upeneus pori – Decapods (Leptochela pugnax), polychaetes, other 80, 17, 3 Demersal 3.51 0.55 19
176. Chromis chromis – Fish, appendicularians, copepods, other 76, 16, 6, 2 Reef-associated 3.18 0.31 25
177. Chromis chromis – Copepods, polychaetes, amphipods, other 67, 10, 9, 14 Reef-associated 4.21 0.72 25
178. Sciaena umbra 13–37 Decapods (Liocarcinus spp., Brachynotus gemmellari), 72, 24, 4 Demersal 3.80 0.64 70
fish, other
179. Umbrina cirrosa 13–53 Decapods (Liocarcinus spp., Upogebia spp.), polychaetes, 78, 9, 5, 8 Demersal 3.51 0.55 70
molluscs, other
180. Euthynnus alletteratus – Fish (Trachurus spp., Scomber spp.) – Pelagic 4.50 0.80 122
181. Sarda sarda – Fish (Sardina pilchardus, Trachurus spp.) – Pelagic 4.50 0.80 91
182. Scomber scombrus – Fish (Sardina pilchardus), crustaceans, gastropods – Pelagic 3.90 0.66 50
183. Thunnus thynnus 28–42 Fish (Sardina pilchardus, Engraulis encrasicolus), 71, 10, 5, 5, 4, 5 Pelagic 4.24 0.72 458
euphausids, cephalopods, stomatopods, decapods, other
184. Thunnus thynnus 49–107 Fish (Engraulis encrasicolus, Sardina pilchardus), 93, 6, 1 Pelagic 4.44 0.78 458
crustaceans, cephalopods
185. Thunnus thynnus 63–129 Fish (Belone acus, Sprattus sprattus), cephalopods (76, 24) Pelagic 4.50 0.74 458
186. Thunnus thynnus – Fish (Sardina pilchardus, Scomber scombrus), fish larvae, – Pelagic 4.00 0.66 458
crustaceans, molluscs
187. Epinephelus aeneus – Fish, cephalopods, crustaceans – Demersal 4.10 0.56 120
188. Epinephelus caninus – Fish, crustaceans, molluscs – Demersal 3.80 0.60 157
189. Epinephelus fasciatus† – Fish, cephalopods – Reef-associated 4.50 0.73 40
190. Epinephelus marginatus 16–98 <30 cm, fish, brachyurans, other 70, 23, 7 Reef-associated 4.13 0.76 150
30–60 cm, fish, brachyurans, molluscs, other 34, 33, 32, 1 3.86 0.65
>60 cm, molluscs, fish, other 73, 19, 8 3.73 0.57
191. Serranus cabrilla – Mysids, decapods, brachyurans, other 48, 39, 12, 1 Reef-associated 3.37 0.42 40
192. Serranus cabrilla Mean SL 13.2 Fish (Gobius niger, Sardina pilchardus), decapods, other 81 (43, 16), 6, 7 Reef-associated 3.30 0.50 40
193. Serranus cabrilla 7–14 Crustaceans, euphausids, decapods ∗∗ Reef-associated 4.37 0.76 40
194. Serranus cabrilla – Fish, cephalopods, crustaceans, polychaetes – Reef-associated 3.80 0.50 40
195. Serranus hepatus – Brachyurans, decapods, amphipods, mysids 57, 32, 6, 5 Demersal 3.47 0.52 25
196. Serranus hepatus Mean SL 7.8 Decapods (Upogebia tipica, Processa nouveli), 29, 29, 24, 15 Demersal 3.50 0.46 25
brachyurans, fish (Boops boops, Gobius sp.), other
197. Serranus hepatus 3–14 Decapods, fish, polychaetes, mysids, amphipods, other 74, 17, 4, 3, 1, 1 Demersal 3.72 0.62 25
198. Serranus hepatus – Crustaceans, decapods, polychaetes, other (63, 13, 11, 13) Demersal 3.73 0.63 25
199. Serranus scriba 10–23 Decapods, fish, brachyurans, other 52, 30, 13, 5 Demersal 3.87 0.66 36
200. Serranus scriba 5–15 Euphausids, decapods, crustaceans ∗∗ Demersal 3.70 0.58 36
201. Siganus luridus 13–25 Algae (Dictyota sp., Cystoseira sp., Sphacelaria sp.) 100 Reef-associated 2.00 0.00 30
202. Siganus luridus 8–30 Red algae (Polysiphonia spp.), brown algae (Sphacelaria 78, 17, 4, 1 Reef-associated 2.00 0.00 30
spp.), green algae, blue-green algae
203. Siganus rivulatus 7–34 Brown algae (Sphacelaria spp.), red algae (Polysiphonia 38, 34, 26, 2 Reef-associated 2.00 0.00 40
spp.), green algae, blue-green algae
204. Boops boops – Algae, copepods, other 49, 48, 3 Demersal 2.53 0.09 36
205. Boops boops – Copepods (Temora stylifera, Isias clavipes), 56, 38, 6 Demersal 3.30 0.47 36
appendicularians, other
 231

Table 1B. Continued

Species Length range Main prey W (or N) Habitat TROPH SE Lmax

206. Dentex dentex 17–83 Fish, cephalopods 74, 26 Benthopelagic 4.50 0.73 100
207. Dentex dentex – Fish, cephalopods – Benthopelagic 4.50 0.73 100
208. Diplodus annularis – Polychaetes, crustaceans, bivalves, other 15, 6, 4, 75 Benthopelagic 3.41 0.35 24
209. Diplodus annularis Mean TL 13.9 Algae, crustaceans, gastropods, polychaetes, other ∗∗ Benthopelagic 3.30 0.45 24
210. Diplodus annularis – Algae, crustaceans, gastropods, polychaetes – Benthopelagic 3.40 0.46 24
211. Diplodus annularis 7–25 Decapods, polychaetes, molluscs, other 31, 23, 23, 23 Benthopelagic 3.32 0.50 24
212. Diplodus puntazzo 28–37 Sponges, algae (Flabellia petiolate, Plocamium 23, 22, 20, 35 Benthopelagic 2.69 0.17 60
cartilagineum), anthozoans, other
213. Diplodus puntazzo Mean TL 3.2 Fish, copepods, amphipods, isopods, algae, other ∗∗ Benthopelagic 3.30 0.48 60
214. Diplodus sargus sargus† 7–40 Bivalves (Mytilus galloprovincialis), echinoderms, algae 58, 19, 10, 13 Demersal 3.04 0.31 45
(Gelidium pusillum, Ulva rigida), other
215. Diplodus sargus sargus† 1–48 Molluscs, fish, decapods, annelids, other 20, 16, 15, 9, 40 Demersal 3.38 0.51 45
216. Diplodus sargus sargus† Mean TL 19.5 Algae (Cymodocea nodosa), bivalves, gastropods, ∗∗ Demersal 3.10 0.40 45
echinoderms, other
217. Diplodus vulgaris 10–24 Bivalves (Mytilus galloprovincialis), ophiurids, 52, 14, 8, 4, 22 Benthopelagic 3.00 0.31 45
polychaetes, algae (Falkenbergia rufolanosa), other
218. Diplodus vulgaris – Polychaetes, echinoderms, amphipods, other 78, 11, 6, 5 Benthopelagic 3.13 0.30 45
219. Diplodus vulgaris 3–40 Fish, echinoderms, annelids, molluscs, other 29, 23, 16, 11, 21 Benthopelagic 3.70 0.55 45
220. Diplodus vulgaris – Tanaidaceans, polychaetes, amphipods, other ∗∗ Benthopelagic 3.20 0.38 45
221. Diplodus vulgaris 13–34 Decapods, echinoderms, bivalves, polychaetes, ∗∗ Benthopelagic 3.50 0.46 45
gastropods, other
222. Lithognathus mormyrus Mean TL 20.1 Bivalves, gastropods, echinoderms, decapods, ∗∗ Benthopelagic 3.30 0.48 55
polychaetes, other
223. Lithognathus mormyrus – Amphipods, decapods, molluscs, gastropods, fish, other – Benthopelagic 3.50 0.52 55
224. Oblada melanura 8–14 Copepods, amphipods, ostracods, other (81, 14, 1, 4) Benthopelagic 3.10 0.20 34
225. Pagellus acarne 8–22 0–12 cm, decapods, fish, polychaetes, molluscs, 45, 19, 14, 7, 6, 9 Benthopelagic 3.65 0.58 36
amphipods, other
12–15 cm, decapods, polychaetes, amphipods, ophiurids, 63, 9, 5, 4, 19 3.54 0.54
other
15–17 cm, decapods, ophiurids, fish, other 65, 26, 4, 5 3.56 0.54
>17 cm, ophiurids, decapods, polychaetes, other 64, 33, 1, 2 3.47 0.45
226. Pagellus bogaraveo 8–16 8–11 cm, eggs, amphipods, bivalves, other (65, 9, 8, 18) Benthopelagic 3.40 0.48 70
11–14 cm, crustaceans, decapods, copepods, fish, other (36, 14, 14, 13, 23) 3.50 0.49
>14 cm, decapods, fish, polychaetes, crustaceans, (24, 16, 15, 15, 15, 15) 3.50 0.50
copepods, other
227. Pagellus erythrinus – Crustaceans, polychaetes, cephalopods, fish, ophiurids, 38, 31, 11, 8, 4, 8 Benthopelagic 3.55 0.46 60
other
228. Pagellus erythrinus 4–47 4–10 cm, amphipods (Caprella sp.), polychaetes, 69, 14, 3, 14 Benthopelagic 3.25 0.48 60
decapods, other
10–15 cm, polychaetes (Sternaspis scutate, Flabelligera 31, 31, 16, 22 3.31 0.46
sp.), amphipods, decapods, other
15–20 cm, polychaetes (Sternaspis scutate, Flabelligera 38, 25, 22, 25 3.37 0.44
sp.), decapods, molluscs, other
>20 cm, decapods, polychaetes, echinoderms, molluscs, 22, 18, 18, 17, 16, 9 3.41 0.47
amphipods, other
229. Pagellus erythrinus 4–45 Annelids, molluscs, decapods, other 42, 21, 20, 17 Benthopelagic 3.36 0.42 60
230. Pagellus erythrinus 3–36 0–5 cm, mysids, crustaceans, bivalves, polychaetes 78, 11, 6, 5 Benthopelagic 3.22 0.40 60
5–10 cm, mysids, crustaceans, polychaetes, decapods, fish, 27, 27, 25, 7, 7, 7 3.39 0.46
cephalopods
10–15 cm, polychaetes, crustaceans, mysids, decapods, 29, 25, 13, 11, 10, 12 3.36 0.44
bivalves, other
15–20 cm, polychaetes, decapods, crustaceans, bivalves, 33, 25, 16, 8, 8, 10 3.43 0.45
ophiurids, other
20–25 cm, polychaetes, crustaceans, ophiurids, decapods, 34, 29, 14, 9, 9, 5 3.37 0.40
bivalves, cephalopods
231. Pagellus erythrinus 7–31 Decapods, fish, gastropods, polychaetes, bivalves, (40, 39, 8, 6, 2, 2, 3) Benthopelagic 3.63 0.53 60
cephalopods, other
232. Pagellus erythrinus – Fish, polychaetes, crustaceans, cephalopods, gastropods, 24, 23, 13, 12, 6, 22 Benthopelagic 3.83 0.53 60
other
233. Pagellus erythrinus – Polychaetes (Sternaspis scutate, Nephthys hystricis), 23, 20, 14, 12, 31 Benthopelagic 3.30 0.45 60
bivalves, decapods, isopods, other
234. Pagellus erythrinus – Decapods (Galathea nexa), polychaetes, gastropods, 25, 24, 20, 7, 24 Benthopelagic 3.16 0.46 60
isopods, other
235. Pagellus erythrinus 9–26 Brachyurans, decapods, polychaetes, other 45, 10, 8, 37 Benthopelagic 3.15 0.48 60
232
Table 1B. Continued

Species Length range Main prey W (or N) Habitat TROPH SE Lmax

236. Pagellus erythrinus 4–22 Brachyurans, decapods, cephalopods, fish, other 34, 6, 5, 4, 51 Benthopelagic 3.12 0.47 60
237. Pagellus erythrinus 6–21 Brachyurans, decapods, cephalopods, polychaetes, other 25, 10, 7, 5, 53 Benthopelagic 3.08 0.44 60
238. Pagellus erythrinus – Crustaceans, polychaetes, fish – Benthopelagic 3.60 0.54 60
239. Pagrus auriga 15–65 Crustaceans, echinoderms, molluscs, other 42, 30, 20, 8 Benthopelagic 3.31 0.48 80
240. Pagrus caeruleostictus 8–40 Molluscs, fish, crustaceans, polychaetes, other 28, 28, 17, 11, 16 Benthopelagic 3.51 0.59 90
241. Pagrus pagrus 13–45 Fish, molluscs, crustaceans, polychaetes, other 52, 22, 10, 6, 10 Reef–associated 3.88 0.67 91
242. Pagrus pagrus 16–49 <21 cm, fish, decapods, echinoderms, other 43, 19,11, 27 Reef-associated 3.74 0.60 91
21–30 cm, fish, echinoderms, brachyurans, anthozoans, 31,17, 14, 13, 8, 17 3.76 0.60
decapods, other
>30 cm, brachyurans, fish, anthozoans, echinoderms, 38, 20, 11, 9, 9, 13 3.61 0.60
decapods, other
243. Pagrus pagrus 4–19 <14 cm, decapods (Upogebia tipica), polychaetes, fish, 53, 25, 19, 3 Reef-associated 3.67 0.60 91
other
>14 cm, decapods (Upogebia tipica), fish, polychaetes 70, 27, 3 3.80 0.65
244. Pagrus pagrus – Decapods, polychaetes (Aponuphis bilineata, Terebellides 54, 21, 18, 7 Reef-associated 3.63 0.56 91
stroemi), fish, other
245. Pagrus pagrus – Fish, decapods, cephalopods, molluscs – Reef-associated 3.90 0.55 91
246. Sarpa salpa 7–37 Algae, epiphytes 96, 4 Benthopelagic 2.06 0.07 51
247. Sarpa salpa 7–37 Algae, epiphytes 99, 1 Benthopelagic 2.02 0.04 51
248. Sarpa salpa 7–37 Algae, epiphytes 96, 4 Benthopelagic 2.06 0.07 51
249. Sarpa salpa Mean TL 3.7 Algae, copepods, hydrozoans, ostracods, amphipods, ∗∗ Benthopelagic 2.50 0.17 51
isopods, other
250. Sarpa salpa 12–24 Rhodophyta (Hypnea musciformis, Gelidium crinale), 49, 27, 24 Benthopelagic 2.00 0.00 51
Phaeophyta, Chlorophyta
251. Sparus auratus† 2–29 Molluscs (Ensis sp.), decapods, annelids, other 60, 16, 13, 11 Demersal 3.42 0.56 70
252. Spondyliosoma cantharus – Mysids, crustaceans 70, 30 Benthopelagic 3.29 0.56 60
253. Lepidopus caudatus 20–69 20–29 cm, euphausids (Meganyctiphanes norvegica, 100 Benthopelagic 3.20 0.40 205
Nyctiphanes couchii)
30–39 cm, euphausids (Meganyctiphanes norvegica), 38, 35, 27 3.69 0.60
decapods, fish
40–49 cm, decapods, fish, other 72, 27, 1 3.84 0.25
50–59 cm, decapods (Pasiphaea sivado), fish, other 69, 25, 6 3.80 0.64
60–69 cm, decapods (Pasiphaea sivado), fish, other 67, 23, 10 3.77 0.63
254. Tripterygion delaisi† Mean TL 6.0 Eggs, copepods, decapods, polychaetes, other (48, 18, 16, 4, 14) Demersal 3.50 0.56 9
255. Tripterygion delaisi† 2–8 Brachyurans, decapods, gastropods, pycnogonids, other (54, 21, 10, 4, 11) Demersal 3.50 0.57 9
256. Uranoscopus scaber – Fish (Callionymus maculatus, Trachurus trachurus), (62, 22, 11, 5) Demersal 3.80 0.57 40
decapods, cephalopods, other
257. Uranoscopus scaber – Fish (Atherina boyeri, Mullus surmuletus), cephalopods, – Demersal 4.00 0.61 40
crustaceans
258. Xiphias gladius – Fish (Sardina pilchardus, Sardinella aurita, Engraulis (100) Pelagic 4.50 0.80 450
encrasicolus)
259. Xiphias gladius 78–192 Cephalopods (Todarodes sagittatus, Ommastrephes 68, 31, 1 Pelagic 4.49 0.54 450
bartramii), fish, decapods
260. Arnoglossus laterna – Amphipods, decapods, polychaetes, other ∗∗ Demersal 3.50 0.49 20
261. Arnoglossus thori – Amphipods 100 Demersal 3.29 0.53 18
262. Bothus podas – Decapods, amphipods, polychaetes 49, 40, 11 Demersal 3.41 0.54 45
263. Bothus podas 6–16 Polychaetes (Nephthys hombergi, Phyllodoce lineata), ∗∗ Demersal 3.40 0.47 45
mysids, cumaceans, amphipods, gastropods, other
264. Citharus linguatula – Fish, decapods – Demersal 3.80 0.65 30
265. Symphurus nigrescens 3–15 3–9 cm, polychaetes, ophiurids, decapods, amphipods, 33,29, 23, 7, 8 Demersal 3.32 0.41 12
other
10–15 cm, decapods, ophiurids, polychaetes, other 47, 22, 21, 10 3.42 0.47
266. Symphurus nigrescens – Polychaetes, decapods, ophiurids, other 39, 26, 17, 18 Demersal 3.20 0.34 12
267. Lepidorhombus boscii 4–32 4–9 cm, decapods (Solenocera membranacea, Processa 66, 15, 6, 13 Demersal 3.54 0.56 40
mediterranea), mysids, fish, other
10–19 cm, decapods (Alpheus glaber, Processa 74, 23, 3 3.80 0.64
mediterranea), fish, other
20–32 cm, decapods (Processa mediterranea, Alpheus 87, 10, 3 3.69 0.61
glaber), fish, other
268. Lepidorhombus boscii 8–20 <12 cm, decapods, mysids, amphipods, other 76, 10, 2, 12 Demersal 3.53 0.56 40
12–16 cm, decapods, fish, mysids, other 74, 20, 2, 4 3.77 0.63
>16 cm, decapods, fish, crustaceans, other 85, 10, 3, 2 3.68 0.61
269. Lepidorhombus boscii 5–29 Crustaceans, decapods, isopods, fish, cephalopods (51, 28, 10, 8, 2) Demersal 3.80 0.61 40
270. Lepidorhombus boscii 5–25 5–10 cm, mysids, decapods, amphipods, other 56, 16, 13, 15 Demersal 3.22 0.48 40
10–20 cm, decapods (Alpheus glaber), mysids, fish, other 48, 26, 14, 12 3.65 0.57
20–25 cm, decapods (Alpheus glaber), fish, other 60, 26, 14 3.85 0.65
 233

Table 1B. Continued

Species Length range Main prey W (or N) Habitat TROPH SE Lmax

271. Lepidorhombus boscii 4–31 Mysids, decapods, isopods, amphipods, fish, brachyurans (35, 23, 6, 6, 5, 5, 20) Demersal 3.50 0.50 40
other
272. Lepidorhombus whiffiagonis 8–20 <12 cm, fish, decapods, mysids, isopods, other 51, 28, 3, 1, 17 Bathydemersal 4.01 0.68 60
12–16 cm, fish, decapods, other 53, 42, 5 4.07 0.70
>16 cm, fish, decapods 71, 29 4.24 0.75
273. Buglossidium luteum† – Amphipods, polychaetes, bivalves ∗∗ Demersal 3.20 0.38 15
274. Solea solea 13–42 13–20 cm, polychaetes, algae ∗∗ Demersal 3.20 0.38 70
20–25 cm, cnidarians, polychaetes, molluscs, decapods, ∗∗ 3.10 0.42
algae
25–30 cm, cnidarians, polychaetes, molluscs, decapods, ∗∗ 3.30 0.46
echinoderms, algae, fish
30–35 cm, cnidarians, polychaetes, molluscs, decapods, ∗∗ 3.30 0.44
echinoderms, algae, fish
35–42 cm, polychaetes, molluscs, decapods, echinoderms, ∗∗ 3.20 0.41
fish, algae
275. Solea solea – Polychaetes, amphipods, tanaidaceans, decapods, 34, 30, 14, 11, 9, 2 Demersal 3.30 0.47 70
bivalves, gastropods
276. Dasyatis chrysonata marmorata† – Fish, polychaetes, gastropods, cephalopods, crustaceans, ∗∗ Demersal 3.70 0.49 60
other
277. Raja miraletus 18–50 Fish, crustaceans, molluscs, polychaetes, other ∗∗ Demersal 3.90 0.59 60
278. Raja radula – Crustaceans, fish, molluscs – Demersal 3.90 0.61 70
279. Rhinobatos rhinobatos 27–180 Decapods, fish, cephalopods 48, 45, 6 Demersal 4.05 0.69 100
280. Scorpaena notata – Fish, appendicularians, copepods, other 80, 10, 5, 5 Demersal 3.49 0.57 24
281. Scorpaena notata – Brachyurans, decapods, other 61, 31, 8 Demersal 3.50 0.57 24
282. Scorpaena notata 2–18 Brachyurans, decapods, amphipods, fish, other 49, 23, 9, 8, 11 Demersal 3.44 0.52 24
283. Scorpaena porcus – Fish, appendicularians, copepods, other 80, 10, 5, 5 Demersal 3.44 0.52 30
284. Scorpaena porcus – Fish, brachyurans, decapods, other 54, 36, 7, 3 Demersal 4.04 0.70 30
285. Scorpaena porcus 2–24 Brachyurans, fish, decapods, other 63, 25, 6, 6 Demersal 3.74 0.64 30
286. Scorpaena porcus 7–20 Fish, crustaceans, other 53, 45, 2 Demersal 4.03 0.70 30
287. Scorpaena porcus 8–23 Brachyurans, fish, decapods, other 42, 29, 20, 9 Demersal 3.82 0.66 30
288. Scorpaena porcus 8–30 Brachyurans, fish, decapods, stomatopods, other 43, 39, 12, 3, 3 Demersal 3.89 0.68 30
289. Scorpaena porcus 7–24 Decapods, crustaceans, bracyurans ∗∗ Demersal 3.40 0.47 30
290. Scorpaena porcus – Brachyurans (Xantho spp., Portunus spp.), decapods, fish – Demersal 3.80 0.66 30
291. Scorpaena scrofa – Fish, decapods 63, 37 Demersal 4.13 0.67 50
292. Scorpaena scrofa 11–23 Fish, brachyurans, decapods, cephalopods 71, 16, 8, 5 Demersal 4.26 0.73 50
293. Scorpaena scrofa 10–25 Fish, crustaceans, other 87, 11, 2 Demersal 4.38 0.77 50
294. Scorpaena scrofa – Fish (Spicara smaris), crustaceans, molluscs – Demersal 4.00 0.66 50
295. Helicolenus dactylopterus 4–29 4–9 cm, fish (Gadiculus argenteus argenteus), decapods, 52, 34, 14 Demersal 4.01 0.69 45
other
10–19 cm, decapods (Alpheus glaber), fish, other 53, 39, 8 3.93 0.67
20–29 cm, decapods (Goneplax rhomboides, Calocaris 81, 9, 8, 2 3.63 0.58
macandreae), tunicates, fish, other
296. Helicolenus dactylopterus 9–17 Decapods, fish, mysids, cephalopods, other (30, 23, 14, 12, 21) Demersal 3.80 0.62 45
297. Chelidonichthys cuculus† – Decapods (Alpheus glaber, Macropipus pusillus), fish, 43, 42, 11, 4 Demersal 3.82 0.61 50
mysids, other
298. Chelidonichthys cuculus† 8–20 Mysids (Leptomysis gracilis, Lophogaster typicus), (78, 7, 7, 5, 3) Demersal 3.60 0.48 50
decapods, cephalopods, fish, other
299. Chelidonichthys gurnardus† 6–39 Decapods (Alpheus glaber, Goneplax rhomboides), 33, 30, 17, 15, 5 Demersal 3.58 0.56 60
mysids, fish, euphausids, other
300. Chelidonichthys lastoviza† – Brachyurans 100 Demersal 3.50 0.60 40
301. Chelidonichthys lastoviza† – Decapods, mysids, other 91, 2, 7 Demersal 3.54 0.53 40
302. Chelidonichthys lastoviza† – Decapods (Liocarcinus maculatus, Liocarcinus arcuatus), (91, 2, 7) Demersal 3.58 0.57 40
mysids, other
303. Chelidonichthys lucerna† 12–35 Decapods (Goneplax rhomboides, Philocheras (55, 14, 12, 10, 9) Demersal 3.40 0.47 75
bispinosus), mysids, fish, molluscs, other
304. Chelidonichthys lucerna† 6–25 Mysids (Gastrosaccus sanctus), decapods, fish, (42, 34, 14, 2, 2, 6) Demersal 3.60 0.54 75
cephalopods, bivalves, other
305. Chelidonichthys lucerna† 6–27 Decapods (Philocheras monacanthus), amphipods, ∗∗ Demersal 3.70 0.48 75
crustaceans, fish, other
306. Chelidonichthys obscurus† 8–23 Decapods (Macropipus depurator, Philocheras sculptus), 53, 21, 14, 12 Demersal 3.73 0.52 20
cephalopods, mysids, other
307. Chelidonichthys obscurus† 8–20 Mysids (Gastrosaccus normani, Anchialina agilis), (78, 17, 5) Demersal 3.50 0.46 20
decapods, other
308. Chelidonichthys obscurus† 11–18 Mysids (Gastrosaccus sp., Lophogaster typicus), (49, 42, 4, 3, 1, 1) Demersal 3.40 0.49 20
decapods, amphipods, copepods, fish, bivalves
234
Table 1B. Continued

Species Length range Main prey W (or N) Habitat TROPH SE Lmax

309. Lepidotrigla cavillone – Mysids (Lophogaster typicus, Anchialina agilis), 74, 18, 8 Demersal 3.29 0.46 20
decapods, other
310. Lepidotrigla cavillone 4–10 Mysids (Gastrosaccus sp., Anchialina agilis), amphipods, (69, 15, 15, 1) Demersal 3.30 0.44 20
decapods, copepods
311. Lepidotrigla cavillone – Mysids (Lophogaster typicus, Paramysis helleri), 76, 10, 14 Demersal 3.17 0.42 20
decapods, other
312. Lepidotrigla cavillone – Decapods (Processa canaliculata), mysids, other 60, 14, 26 Demersal 3.10 0.41 20
313. Lepidotrigla cavillone – Mysids (Paramysis helleri, Lophogaster typicus), 71, 16, 13 Demersal 3.37 0.48 20
decapods, other
314. Lepidotrigla cavillone – Mysids (Lophogaster typicus, Paramysis helleri), 75, 10, 15 Demersal 3.28 0.44 20
decapods, other
315. Lepidotrigla cavillone – Decapods (Upogebia tipica, Solenocera membranacea), 55, 42, 3 Demersal 3.43 0.52 20
mysids, crustaceans
316. Trigla lyra 4–39 4–9 cm, ophiurids, hydrozoans, other 85, 12, 3 Demersal 3.48 0.35 60
10–19 cm, ophiurids, decapods, other 77, 21, 2 3.48 0.42
20–29 cm, decapods (Goneplax rhomboides, Pontocaris 66, 34 3.53 0.52
lacazei), ophiurids
30–39 cm, decapods, ophiurids 65, 35 3.53 0.52
317. Trigla lyra 23–50 Crustaceans (Anapagurus laevis, Monodaeus couchi), (50, 39, 11) Demersal 3.70 0.52 60
polychaetes, echinoderms
318. Trigla lyra 6–30 <18 cm, mysids (Lophogaster typicus), decapods, other 68, 18, 14 Demersal 3.31 0.47 60
>18 cm, mysids (Lophogaster typicus), brachyurans, 48, 24, 20, 8 3.38 0.50
decapods, other
319. Trigla lyra 5–33 <18 cm, mysids (Lophogaster typicus), decapods, other 74, 13, 13 Demersal 3.28 0.45 60
>18 cm, mysids (Lophogaster typicus), brachyurans, 46, 27, 16, 11 3.38 0.51
decapods, other
320. Centroscymnus coelolepis 19–75 19–39 cm, cephalopods, fish, decapods, other 62, 20, 5, 13 Bathydemersal 4.16 0.48 120
40–75 cm, cephalopods, fish, other 88, 6, 6 4.35 0.40
321. Dalatias licha† 32–101 Fish (Trachyrhynchus trachyrhynchus, Notoscopelus 75, 15, 5, 5 Bathydemersal 4.35 0.75 182
elongatus), decapods, molluscs, other
322. Etmopterus spinax 10–49 Fish (Engraulis encrasicolus, Notolepis rissoi), 56, 26, 16, 2 Bathydemersal 4.33 0.67 60
cephalopods, decapods, other
323. Etmopterus spinax – Fish (Engraulis encrasicolus, Notolepis rissoi), 49, 19, 11, 21 Bathydemersal 3.80 0.58 60
crustaceans, molluscs, other
324. Hippocampus hippocampus – Amphipods, copepods 53, 47 Demersal 3.15 0.39 15
325. Syngnathus acus – Brachyurans, polychaetes, decapods, fish, other 64, 17, 7, 4, 8 Demersal 3.44 0.40 46
326. Syngnathus acus – Decapods, amphipods, mysids, other 75, 16, 8, 1 Demersal 3.47 0.55 46
327. Syngnathus typhle – Fish, mysids, decapods 84, 11, 5 Demersal 4.31 0.75 30
328. Torpedo marmorata 11–62 <20 cm, fish, cephalopods, decapods, polychaetes 75, 20, 3, 2 Demersal 4.45 0.72 60
20–40 cm, fish, cephalopods 95, 5 4.50 0.78
>40 cm, fish 100 4.50 0.80
329. Torpedo torpedo 12–43 <20 cm, fish, decapods, cephalopods 74, 18, 8 Demersal 4.32 0.74 60
20–30 cm, fish, decapods 95, 5 4.45 0.79
>30 cm, fish 100 4.50 0.80
330. Capros aper 3–15 3–9 cm, euphausids, euphausids larvae, copepods, other 51, 25, 21, 3 Demersal 3.16 0.36 30
10–15 cm, euphausids, copepods, other 54, 22, 24 3.21 0.38
331. Zeus faber – Fish 100 Benthopelagic 4.50 0.80 90
332. Zeus faber 5–51 <14 cm, fish (Cepola macrophthalma, Deltentosteus 89 (39, 33), 9, 2 Benthopelagic 4.36 0.77 90
quadrimaculatus), mysids, other
>14 cm, fish (Cepola macrophthalma, Spicara flexuosa), 99.7 (65, 19), 0.3 4.50 0.80
other

reported length classes, Li , was quantified using the
following equation (Cortés, 1999):
TROPHLi = TROPHL∞ (1 − e−KLi ),
where TROPHL∞ is the asymptotic TROPH and K
is the rate at which TROPHL∞ is approached, with
both estimated using Marquardt’s (1963) algorithm for
fitting nonlinear least-squares regression. An F test
of significance of the entire nonlinear regression was
obtained from the (regression mean square)/(residual
mean square) ratio, which was compared to the critical
value F0.5(1),v1,v2, where v1 = regression degrees of
freedom (df) = 2 and v2 = residual df = (total number
of length classes) = 3 (Zar, 1999).
Results
Overall, 332 datasets of diet composition were
gathered, which corresponded to 146 fish species, 59

families and 21 orders (Table 1A). The maximum
reported body size (Lmax ) of the 146 species ranged
from 4 cm, for Apletodon dentatus, to 458 cm, for
Thunnus thynnus (Table 1B).
The majority of the datasets referred to the Western
Mediterranean (183 datasets, 83 of which referred to
the Gulf of Lions, including the Gulf of Marseille,
and 65 to the Balearic Sea) and, to a lesser extent,
to the Eastern Mediterranean (102 datasets, 85 of
which referred to the Aegean Sea) (Figure 1). Only 47
datasets were derived from the Central Mediterranean
Sea (Figure 1). Furthermore, only 33 datasets were
derived from the southern rim of the Mediterranean
Sea (Figure 1).
The compiled datasets were based on samples
collected mainly with trawls (Figure 2a) and on a
monthly or seasonal basis (Figure 2b). The sampling
gear (Figure 2a) and sampling frequency (Figure 2b)
was not reported for 60 and 76 datasets, respectively.
Diet composition was expressed as percentages of
numerical abundance (N), gravimetric abundance (W),
frequency of occurrence (F) or rank relative abundance
(RA) of prey in relation to the number of stomachs
containing food (Figure 2c). For 222 datasets, the
stomach content was expressed using W along with
N and/or F whereas for the remaining 110 datasets
it was expressed using N and/or F or RA (Figure
2c). The length range of the specimens studied was
provided for 169 datasets and the mean length for
26 datasets, whereas no information on the length
structure of the specimens studied was available for
the remaining 137 datasets (Figure 2d). Finally, the
number of empty stomachs was provided for 213
datasets (64.2%) (Table 1A).
The total number of stomachs analyzed per dataset,
which ranged from 1 for eight datasets to 5,223 for
Saurida undosquamis in SE Mediterranean in 1980–
1981, was not reported for 35 datasets (Figure 3).
For about half of the datasets (156, 47%) the total
number of stomachs analyzed was lower than 200
(Figure 3).
The estimated TROPH values (Table 1B) ranged
from 2.0 to 4.5 units and displayed three main
modes, at 3.4–3.5, 3.8–3.9 and 4.3–4.4 units, and two
minor ones at 2.0–2.1 and 2.5–2.6, and three normal
distribution components were identified (Figure 4).
The above-mentioned modes and normal distributions
might correspond to the following general functional
trophic groups:
(a) Pure herbivores (TROPH = 2.0–2.1, mean
= 2.02, SD = 0.03), which were very rare among
235
the datasets reviewed here (Figure 4). They were
represented by the reef-associated Siganus luridus
and Siganus rivulatus datasets (in the Dodecanese
in 1985–1986 and in SE Mediterranean, undated)
and the benthopelagic Sarpa salpa (in the Southern
Adriatic Sea in 1989; and in the Gulf of Lions, Gulf
of Marseille and Corsican waters, all undated). The
above-mentioned stocks all feed on red, brown, green
and blue-green algae, with the last three Sarpa salpa
stocks also accidentally consuming small quantities of
epiphytes (Table 1B).
(b) Omnivores with a preference for vegetable
material (2.1 < TROPH < 2.9, mean = 0.50, SD =
0.12), but also feeding on other prey, such as sponges,
hydrozoans, anthozoans, polychaetes, ostracods,
isopods, amphipods and copepods. They were very
rare among the datasets reviewed here and represented
by the following species (Table 1B): Gobius bucchichi
in the Northern Adriatic Sea (undated); Boops boops
in the Gulf of Marseille in 1980; Diplodus puntazzo in
the Western Mediterranean Sea in 1992-1993; Sarpa
salpa in Western Sicilian waters in 1991; and 13–20
cm Solea solea in the Western Mediterranean Sea in
1976–1978.
(c) Omnivores with a preference for animals (2.9
< TROPH < 3.7, mean = 3.40, SD = 0.19) feeding
on a wide variety of prey (e.g., algae, foraminifera,
brachyurans, balanoids, ostracods, bivalves, ascid-
ians, amphipods, appendicularians, annelids, isopods,
gastropods, cumaceans, cnidarians, ophiurids, poly-
chaetes, cladocerans, mysids, euphausids, fish
larvae, cephalopods) (Figure 4). They were the most
numerous group among the datasets reviewed (Figure
4). They were mainly represented by the species of
the families Blenniidae, Bothidae, Centracanthidae,
Gobiidae, Labridae, Lotidae, Macrouridae, Mullidae,
Ophidiidae, Soleidae, Triglidae (Figure 5), Cyno-
glossidae (Symphurus nigrescens), Alepocephal-
idae (Alepocephalus rostratus), Clupeidae (Sardina
pilchardus) and Engraulidae (Engraulis encrasicolus)
(Table 2). Although the members of this group
occurred in all habitat types, they were mainly distrib-
uted in the bathypelagic, benthopelagic and demersal
habitats (Figure 6).
(d) Carnivores with a preference for large deca-
pods, cephalopods and fish (3.7 < TROPH <
4.5), which were the next most abundant group
among the datasets reviewed (Figure 4). They were
mainly represented by the species of the families
Dalatiidae, Lophiidae, Scombridae, Scyliorhinidae,
Synodontidae, Torpedinidae (Figure 5), Merlucciidae
236
Figure 1. Spatial allocation of the 332 compiled datasets on diet composition of Mediterranean fish as well as names of main seas, gulfs and
islands. Division of Mediterranean Sea into Western, Central and Eastern parts follows the system of the General Fisheries Council for the
Mediterranean Sea (GFCM).
(Merluccius merluccius), Sebastidae (Helicolenus
dactylopterus), Xiphiidae (Xiphias gladius) and
Zeidae (Zeus faber) (Table 2). Although the members
of this group also generally occurred in all habitat
types, they were mainly distributed in the bathyde-
mersal, pelagic and reef-associated habitats (Figure 6).
This group was further divided into two subgroups:
one exhibiting a preference for decapods and fish,
feeding also on polychaetes, brachyurans, echino-
derms, crustaceans, molluscs, mysids and cephalo-
pods (3.7 < TROPH < 4.0, mean = 3.85, SD = 0.09)
and another one exhibiting a preference for fish and
cephalopods (4.0 < TROPH < 4.5, mean = 4.38, SD
= 0.12), feeding also on crustaceans and decapods.
Among the latter subgroup, the species characterized
by datasets with the highest TROPH (4.5; Dentex
dentex, Molva macrophthalma, Seriola dumerili,
Xiphias gladius, Thunnus thynnus, Zeus faber,
Torpedo marmorata, Torpedo torpedo, Epinephelus
fasciatus, Euthynnus alletteratus and Sarda sarda)
and with TROPH values ranging between 4.4 and
4.5 (Saurida undosquamis, Seriola dumerili, Conger
conger, Merluccius merluccius, Thunnus thynnus and
Xiphias gladius) (Table 1B, Table 2) can be considered
as top carnivores when compared to the remaining
species reviewed here.
The family Sparidae, which was characterized
by the highest number of species and datasets, was
represented in all functional trophic groups (Figure
5). In addition, the families Carangidae, Congridae,
Gadidae, Scorpaenidae, Serranidae, Scophthalmidae
and Syngnathidae all included representatives in both
trophic groups (c) and (d) (Figure 5).
The most intensively studied species were Mullus
barbatus, Merluccius merluccius, Pagellus erythrinus
and Mullus surmuletus, represented by more than
10 datasets each, followed by Scorpaena porcus and
Lepidotrigla cavillone (8 and 7 datasets respectively)
(Table 2). In addition, 69 species were represented by
2 to 5 datasets (Table 2) each and the remaining 71
species by 1 dataset each. The TROPH values esti-
mated from the different datasets of the same species
varied with area (Table 2). The coefficient of variation
ranged between 0.0 and 20% and for most species (49,
65%) it was smaller than 5% whereas for 6 species
it was higher than 10% (Table 2). The TROPH differ-
ence (i.e., difference between maximum and minimum
TROPH per species) ranged between 0.0 and 1.3 units,
 237

Figure 2. (a) Sampling gear (for other gears see Table 1A), (b) frequency of sampling, (c) indices used for the expression of diet composition
data (W = gravimetric contribution, N = numerical contribution, F = frequency of occurrence of prey, RA=rank relative abundance) and (d)
type of length of examined specimens reported, for the 332 datasets on diet composition of Mediterranean fish compiled in the present study.
NR = Not reported.

depending on species (Figure 7). For most species (43,
57%) TROPH differences ranged between 0.0 and 0.3
units whereas for 16 species they were greater than 0.5
units (Figure 7).
Across the datasets analyzed, TROPH and mean
TROPH per species increased with a corresponding
increase in species’ Lmax (Figure 8). The latter
accounted for 20–23% of the variance in TROPH
and this percentage increased to 32–36% when few
outliers, corresponding to all datasets of the pure
herbivores Siganus luridus, Siganus rivulatus and
Sarpa salpa, were excluded from the analysis (Figure
8).
Stomach content data for different length classes
of the same stock were available for 38 datasets only
(Table 3). TROPH increased between the smallest
and largest length classes for 32 datasets (Table 3).
TROPH differences ranged from −0.4 to 1.1 units,
with about half of the datasets (i.e., 17, 45%) exhibit-
ing a difference of 0.0–0.2 units (Figure 9).
The relationship between TROPH and the mid-
point of each length class was estimated only for 11
datasets for which data were available for more than
three length classes (Table 3). The asymptotic model
provided a significant (i.e., regression F value > crit-
ical value F0.5(1),v1,v2) fit for 9 datasets (Figure 10,
Table 4). It described the datasets adequately (i.e., R2
> 0.73), with two exceptions (Figure 10b, c) for which
R2 was 0.40 and 0.42 (Table 4). For these two data-
sets, TROPH increased linearly with length (Chimaera
monstrosa: TROPHLi = 3.36 + 0.034Li, R2 = 0.90, P
< 0.05; Caelorinchus caelorhincus: TROPHLi = 2.97
+ 0.54Li, R2 = 0.93, P < 0.05). The asymptotic model
did not provide a fit for two datasets (Figure 10a, f) for
which TROPH did not change significantly (P > 0.05)
with length.
238

Figure 3. Histogram of the total number of stomachs on which the 332 datasets on diet composition of Mediterranean fish were based. NR =
Not reported.

Figure 4. Histogram of all TROPH values of Mediterranean fish listed in Table 1B, including those referring to different length classes (N =
416), together with the three normal distributions identified. The latter are characterized by their means (left) and standard deviations (right, in
parentheses). The two classes at the left (i.e., midpoint 2.05) and right (i.e., midpoint 4.55; all cases had TROPHs = 4.5) edges of the TROPH
distribution were not considered in the analysis.
 239

Figure 5. Box – Whisker plots of the TROPH values of Mediterranean fish by major family. The central box covers 50% of the data, the
whiskers extend out to the minimum and maximum values of the data, the vertical line within the box is the median and the black dot is the
mean. Numbers in parentheses indicate number of species (left) and number of datasets (right) per family. The vertical lines indicate the trophic
groups identified in Figure 4.

Discussion abundant yet important components of the Mediter-

From Tables 1 and 2 it became apparent that stomach
contents, and thus TROPH estimates, are avail-
able for many of the most important, in terms of
reported catches, Mediterranean fish (i.e., Engraulis
encrasicolus, Sardina pilchardus, Merluccius merluc-
cius, Sparus auratus, Trachurus spp., Boops boops,
Mullus spp., Thunnus thynnus, Micromesistius
poutassou: their percentage contribution to the 1996
catch ranged from 29.2 to 1.4% respectively). Yet, we
did not find any data for the feeding habits of many
other commercially important fish, which are recorded
separately in the statistical Bulletins of the Food
and Agriculture Organisation (FAO) and contribute
considerably to the overall reported Mediterranean
catches, such as: (a) Scomber japonicus, Sardinella
aurita, Sprattus sprattus, Merlangius merlangus (all
of which contributed more than 1% of the total 1996
Mediterranean catch); (b) Auxis spp., Pomatomus
saltatrix, Mustelus spp., Psetta maxima, Sphyraena
spp. (all of which contributed more than 0.1% of the
total 1996 Mediterranean catch); and (c) other less
ranean ecosystems (e.g., Belone belone, Dentex
macrophthalmus, Squalus acanthias, Caranx spp.,
Squatina squatina, Platichthys flesus, Katsuwonus
pelamis, Lamna nasus, Thunnus obesus, Istiophorus
albicans, Pleuronectes platessa). Thus, the study of
the feeding habits of the above-mentioned species,
especially of those from the bathypelagic, bathyde-
mersal, reef-associated and pelagic habitats, which
were underrepresented in our compilation (Figure 6),
together with similar information on other organisms
such as large crustaceans and cephalopods, will be
essential for the development of Ecopath models, for
the estimation of “Primary Production Required” to
support the Mediterranean fisheries, as well as for the
evaluation of the ‘fishing down the food web’ effect at
a small, local scale in the Mediterranean Sea, where
such information is lacking to a great extent (Stergiou
and Polunin, 2000).
In marine ecosystems, consumers have TROPH
values that range between 2.0, for herbivorous/
detrivorous organisms, and 5.0, for piscivorous/
carnivorous organisms, the latter being rare even
240
Table 2. Minimum, maximum and mean fractional trophic level (TROPH) values for all Mediterranean
fish species represented by more than one dataset (N). CV: coefficient of variation. When TROPH was
available per length class, the mean of all classes was taken

TROPH
Species N Mimimum Maximum Mean CV (%)

Siganus luridus 2 2.00 2.00 2.00 0.00

Sarpa salpa 5 2.00 2.50 2.13 9.90
Boops boops 2 2.53 3.30 2.92 18.70
Diplodus puntazzo 2 2.69 3.30 3.00 14.40
Spicara smaris 2 3.00 3.10 3.05 2.30
Sardina pilchardus 2 3.10 3.20 3.15 2.30
Spicara maena 3 3.00 3.30 3.15 4.80
Symphodus mediterraneus 2 3.10 3.22 3.16 2.70
Solea solea 2 3.04 3.30 3.17 5.80
Diplodus sargus sargus 3 3.04 3.38 3.17 5.70
Gobius auratus 2 3.10 3.30 3.20 4.40
Symphodus tinca 3 3.14 3.26 3.20 1.90
Symphodus cinereus 3 3.10 3.30 3.23 3.40
Deltentosteus quadrimaculatus 3 3.11 3.30 3.24 3.40
Symphodus ocellatus 4 2.99 3.40 3.24 1.00
Nezumia aequalis 2 3.20 3.29 3.25 2.00
Pomatoschistus bathi 2 3.20 3.30 3.25 2.20
Mullus barbatus 16 2.79 3.57 3.27 6.90
Lepidotrigla cavillone 7 3.10 3.43 3.28 3.40
Caelorinchus caelorhincus 2 3.20 3.36 3.28 3.40
Symphurus nigrescens 2 3.20 3.37 3.29 3.70
Hymenocephalus italicus 2 3.20 3.40 3.30 4.30
Diplodus vulgaris 5 3.00 3.70 3.31 8.70
Gobius niger 3 3.20 3.47 3.32 4.10
Diplodus annularis 4 3.30 3.41 3.36 1.70
Symphodus rostratus 3 3.30 3.41 3.37 1.80
Pagellus erythrinus 12 3.08 3.83 3.37 7.00
Coris julis 4 3.27 3.63 3.38 5.10
Mullus surmuletus 11 3.16 3.58 3.38 4.30
Lithognathus mormyrus 2 3.30 3.50 3.40 4.10
Bothus podas 2 3.40 3.41 3.41 2.40
Gaidropsarus mediterraneus 2 3.41 3.42 3.42 0.20
Engraulis encrasicolus 3 3.40 3.50 3.43 1.70
Trachyrincus scabrus 3 3.20 3.66 3.45 6.80
Gaidropsarus vulgaris 2 3.42 3.49 3.46 1.40
Syngnathus acus 2 3.44 3.47 3.46 0.60
Trigla lyra 4 3.33 3.51 3.47 5.00
Scorpaena notata 3 3.44 3.50 3.48 0.90
Lepidion lepidion 2 3.30 3.67 3.49 7.50
Alepocephalus rostratus 2 3.40 3.59 3.49 3.80
Trachurus mediterraneus 4 3.20 3.70 3.50 6.20
Tripterygion delaisi 2 3.50 3.50 3.50 0.00
Ophidion barbatum 2 3.47 3.56 3.52 1.80
Chelidonichthys lastoviza 3 3.50 3.58 3.54 1.10
Chelidonichthys obscurus 3 3.40 3.73 3.54 4.80
Chelidonichthys lucerna 3 3.40 3.70 3.57 4.30
Serranus hepatus 4 3.47 3.73 3.61 3.90
 241
Table 2. Continued

TROPH
Species N Mimimum Maximum Mean CV (%)

Gaidropsarus biscayensis 2 3.52 3.72 3.62 3.90

Lepidorhombus boscii 5 3.50 3.80 3.64 3.13
Gnathophis mystax 2 3.62 3.67 3.65 0.96
Upeneus moluccensis 2 3.40 3.89 3.65 9.50
Trachurus trachurus 4 3.40 3.90 3.65 5.70
Trisopterus minutus 5 3.39 3.81 3.65 5.30
Chromis chromis 2 3.18 4.21 3.70 19.70
Conger conger 3 3.20 4.49 3.70 18.60
Chelidonichthys cuculus 2 3.60 3.82 3.71 4.20
Serranus cabrilla 4 3.30 4.37 3.71 13.30
Gadiculus argenteus argenteus 2 3.55 3.90 3.73 1.60
Pagrus pagrus 5 3.63 3.90 3.77 3.10
Scorpaena porcus 8 3.40 4.04 3.77 6.40
Serranus scriba 2 3.70 3.87 3.79 3.20
Helicolenus dactylopterus 2 3.80 3.86 3.83 1.10
Uranoscopus scaber 2 3.80 4.00 3.90 3.60
Micromesistius poutassou 5 3.56 4.39 3.92 7.90
Galeus melastomus 3 3.70 4.26 3.97 1.99
Etmopterus spinax 2 3.80 4.33 4.07 9.20
Saurida undosquamis 2 3.80 4.49 4.15 11.80
Lophius budegassa 2 3.90 4.40 4.15 3.00
Merluccius merluccius 13 3.75 4.45 4.17 5.50
Scorpaena scrofa 4 4.00 4.38 4.19 3.90
Seriola dumerili 5 3.83 4.50 4.21 8.00
Thunnus thynnus 4 4.24 4.50 4.30 5.30
Zeus faber 2 4.43 4.50 4.47 1.10
Xiphias gladius 2 4.49 4.50 4.50 0.20
Dentex dentex 2 4.50 4.50 4.50 0.00

in fish such as sharks (Cortés, 1999), character-
izing only specialized predators of marine mammals
(e.g., killer whales or polar bears; Pauly et al.,
1995, 1998c). In the Mediterranean Sea, the highest
TROPH value, 4.5, was recorded for Dentex dentex,
Epinephelus fasciatus, Euthynnus alletteratus, Molva
macrophthalma, Sarda sarda, Seriola dumerili,
Thunnus thynnus, Torpedo marmorata, Torpedo
torpedo, Xiphias gladius and Zeus faber (>14 cm),
followed by Saurida undosquamis, Conger conger and
Merluccius merluccius (TROPH values from 4.4 to
4.49). Such TROPH values are much higher than those
estimated for many sharks, which generally range
between 3.1 and 4.7 (Cortés, 1999), and many marine
mammals, which generally range between 3.2 and 4.5
(Pauly et al., 1995, 1998c), with members of both
groups being generally considered as top predators.
We also estimated TROPH from the weight of the prey
in the stomach contents of one monk seal Monachus
monachus, one bottle-nosed dolphin Tursiops trun-
catus, one dolphin Stenella coeruleoalba and one
leatherback turtle Dermochelys coriacea from the
Mediterranean Sea (Table 5). Our TROPH estimates
(Figure 4) were all lower than those for Tursiops trun-
catus and Monachus monachus, whereas the TROPH
of Stenella coeruleoalba was 4.5 (Table 5). Hence,
the fish species mentioned above, along with marine
mammals, must be considered as top predators in the
Mediterranean Sea.
The analysis of the estimated TROPHs indicated
a number of different functional trophic groups (i.e.,
pure herbivores; omnivores with a preference for vege-
table material; omnivores with a preference for a
variety of animal prey; carnivores with a preference
242

Figure 6. Box – Whisker plots of the fractional trophic level (TROPH) values of Mediterranean fish by habitat type. The central box covers
50% of the data, the whiskers extend out to the minimum and maximum values of the data, the vertical line within the box is the median and the
black dot is the mean. Numbers in parentheses indicate number of datasets per habitat type. Habitat type was extracted from “FishBase online”
(www.fishbase.org). The vertical lines indicate the trophic groups identified in Figure 4.

Figure 7. Frequency distribution of the differences between the minimum and maximum TROPH values for all Mediterranean fish species
represented by more than one dataset (N = 75; see Table 2).
 243

Table 3. Fractional trophic level (TROPH) values for the smallest and largest length class of Mediterranean
fishes for which diet composition was available for different length classes. N: number of length classes

TROPH
Species Area N Smallest Largest
class class

Zeus faber G. Evvoikos 12 3.40 4.50

Merluccius merluccius C Mediterranean 7 3.43 4.20
Chimaera monstrosa W Mediterranean 5 3.38 3.59
Lepidopus caudatus W Mediterranean 5 3.20 3.77
Pagellus erythrinus C Tyrrhenian Sea 5 3.22 3.37
Solea solea W Mediterranean 5 2.30 3.20
Caelorinchus caelorhincus W Mediterranean 4 3.17 3.60
Merluccius merluccius G. Lions 4 3.98 4.43
Pagellus acarne G. Evvoikos 4 3.65 3.47
Pagellus erythrinus G. Lions 4 3.25 3.41
Trigla lyra W Mediterranean 4 3.48 3.53
Epinephelus marginatus G. Annaba 3 4.13 3.73
Gnathophis mystax W Mediterranean 3 3.55 3.60
Helicolenus dactylopterus W Mediterranean 3 4.01 3.63
Lepidorhombus boscii W Mediterranean 3 3.54 3.69
Lepidorhombus boscii G. Valencia 3 3.53 3.68
Lepidorhombus boscii Tyrrhenia Sea 3 3.22 3.85
Lepidorhombus whiffiagonis G. Valencia 3 4.01 4.24
Micromesistius poutassou W Mediterranean 3 3.97 4.04
Nezumia aequalis W Mediterranean 3 3.29 3.09
Pagellus bogaraveo N Aegean Sea 3 3.40 3.50
Pagrus pagrus Dodecanese 3 3.74 3.61
Seriola dumerili Tyrrhenia Sea 3 3.50 4.30
Seriola dumerili G. Castellammare 3 3.40 4.37
Torpedo marmorata Egypt 3 4.45 4.50
Torpedo torpedo Egypt 3 4.32 4.50
Trachyrincus scabrus W Mediterranean 3 3.58 3.83
Alepocephalus rostratus W Mediterranean 2 3.39 3.91
Capros aper W Mediterranean 2 3.16 3.21
Centroscymnus coelolepis W Mediterranean 2 4.16 4.35
Gaidropsarus biscayensis C Adriatic Sea 2 3.40 3.63
Galeus melastomus Catalan Sea 2 3.86 4.18
Micromesistius poutassou G. Patraikos 2 3.34 3.78
Pagrus pagrus Cretan waters 2 3.67 3.80
Seriola dumerili Catalan Sea 2 4.49 4.49
Symphurus nigrescens W Mediterranean 2 3.32 3.42
Trigla lyra G. Saronikos 2 3.31 3.38
Trigla lyra N Aegean Sea 2 3.28 3.38
244

Figure 8. Relationship between maximum reported body length (Lmax , cm) (extracted from “FishBase online”; www.fishbase.org) and: (a)
TROPHs of 330 out of the 332 datasets compiled; and (b) mean TROPH per species. Lmax was not available for Notacanthus bonaparte and
Pomatoschistus quagga. When the lower outliers (corresponding to all datasets of the pure herbivores Siganus spp. and Sarpa salpa) were not
included in the analysis, the regression equations were for (a) Log(TROPH) = 0.433 + 0.075Log(Lmax ) [SE(b) = 0.006; r2 = 0.32, n = 322, P
< 0.01] and for (b) Log(Mean TROPH) = 0.440 + 0.073 Log(Lmax ) [SE(b) = 0.008; r2 = 0.36, n = 141, P < 0.01].

Table 4. Estimated values of the parameters TROPHL∞ and K, together with their standard errors (SE), of the relationship between
factional trophic level (TROPHLi ) and body length (mid-point of reported length classes, Li ; in cm) [TROPHLi = TROPHL∞ (1 −
e−KLi )] for nine fish stocks (see Figure 10). R2 : coefficient of determination. N: number of length classes

Species Area TROPHL∞ SE K SE R2 N

Chimaera monstrosa W Mediterranean 3.50 0.047 0.13 0.027 0.40 5

Caelorinchus caelorhincus W Mediterranean 3.44 0.117 0.35 0.109 0.42 4
Merluccius merluccius G. Lions 4.40 0.034 0.21 0.014 0.95 4
Merluccius merluccius C Mediterranean 4.25 0.117 0.14 0.019 0.83 6
Pagellus erythrinus G. Lions 3.37 0.028 0.45 0.058 0.73 4
Pagellus erythrinus C Tyrrhenian Sea 3.39 0.016 1.20 0.082 0.89 5
Lepidopus caudatus C Mediterranean 3.88 0.079 0.076 0.009 0.89 5
Solea solea W Mediterranean 3.58 0.348 0.075 0.022 0.79 5
Zeus faber G. Evvoikos 4.51 0.048 0.186 0.011 0.92 12
 245

Figure 9. Histogram of the differences between the TROPH values of the smallest and largest length classes for all Mediterranean fish species
for which diet composition was available for different length classes (N = 38; see Table 3).

Table 5. Fractional trophic levels (TROPH) estimated using diet composition data from reported stomach
contents, of four species of marine mammals and turtles in the Mediterranean Sea

Species Area TROPH Reference

Dermochelys coriacea SW Sicilian waters 4.00 Jereb and Ragonese (1990)

Monachus monachus C Aegean Sea 4.57 Cebrian et al. (1990)
Stenella coeruleoalba W Mediterranean Sea 4.50 Duguy et al. (1979)
Tursiops truncatus Livorno waters 4.87 Voliani and Volpi (1990)

for decapods and fish; and carnivores with a prefer-
ence for fish and cephalopods: Figure 4). Although
we are not aware of similar studies in other ecosys-
tems, it would be interesting to see what the patterns of
TROPH distribution are in other regions of the world’s
oceans.
The trophic groups identified (Figure 4) are provi-
sional and subject to revision (in terms of mean
TROPH and range) when more datasets for other
species/habitats (see above) and areas (especially from
the Southern Mediterranean, which was underrepre-
sented in our compilation: Figure 1) become avail-
able. In any case, such a grouping may serve as the
basis for the maintenance of trophic level balance in
the Mediterranean Sea, by evaluating corresponding
indicators (e.g., aggregate annual removal from each
trophic group) and reference points (e.g., maximum
% removal from each trophic group). This will set the
basis for an ecosystem approach to the management of
the Mediterranean Sea.
Only eight datasets belonging to five shark species
(i.e., three datasets of Galeus melastomus; one
of Scyliorhinus canicula; one of Centroscymnus
coelolepis; one of Dalatias licha; and two of
Etmopterus spinax) were included in our review and
their TROPH values were higher than those estimated
by Cortés (1999) (Table 6). This must be attributed
to the fact that Cortés’ (1999) estimates are mean
estimates per species, irrespective of the distribu-
tional area of the species. Indeed, our results showed
that within-species TROPH estimates might differ
considerably spatially (Table 2). Yet, the differences
observed in the present study (Table 2) might also
reflect the effect of factors other than spatial distri-
246
Table 6. Comparison of the fractional trophic levels (TROPH)
of five shark species included in the study of Cortés (1999) with
those estimated in the present study
TROPH
Species Present study Cortés (1999)
Galeus melastomus 3.70–4.26 3.70
Scyliorhinus canicula 3.80 3.60
Centroscymnus coelolepis 4.35 4.20
Dalatias licha 4.35 4.10
Etmopterus spinax 3.80–4.33 3.80
bution, such as different methodologies and investi-
gators, temporal variability in feeding, and differences
in the length structure of specimens analyzed.
Within-species TROPHs also differ with time
(e.g., Pauly and Palomares, 2000). Unfortunately,
our compilation did not include datasets referring
to the same stock for more than 2–3 different time
periods (Table 1). Thus, the identification of temporal
trends in the TROPH of Mediterranean fish was not
possible with the available datasets. This is mainly the
result of the fact that, with few exceptions, research
on this and other fisheries issues in the Mediter-
ranean Sea lacks continuity in sampling, which seri-
ously hampers management (Stergiou, 2000; Lleonart,
2000).
TROPH generally increases ontogenetically with
size for stocks for which such data are available (Pauly
et al., 1998b; Pauly and Palomares, 2000; Polunin and
Pinnegar, 2000). Our analysis of the Mediterranean
datasets also indicated that for most species TROPH
increased with length (Table 3) and this increase was
asymptotic in shape (Figure 10). Such a relationship
between TROPH and length may mask the identifi-
cation of the “fishing down the food web” effect
(Pauly et al., 1998a, b, 2000a). Recently, a method to
circumvent such a masking effect has been developed
by Pauly and Palomares (2000) and Pauly et al.
(2001).
A similar relationship between TROPH and length
is also true for among-species comparisons (Figure
8; see also: Cortés, 1999; Pauly et al., 2001). It
is worthy to point out that both within- and among-
species changes in TROPH with size will not hold
for pure herbivores, such as Siganus spp. and Sarpa
salpa. Indeed, all stocks of these species deviated (i.e.,
outliers) from the general TROPH-Lmax relationships
built (Figure 8).
The among- and within-species asymptotic rela-
tionships between TROPH and length can be
explained in the light of foraging behaviour and prey
selection theory. For most fish, the sizes of prey
consumed generally increase with a corresponding
increase in predator size. This is true for compar-
isons within-species (e.g., Keast and Webb, 1966;
Kaiser and Hughes, 1993; Juanes and Conover, 1994;
Pauly et al., 1998b) and among-species (e.g., Juanes,
1994; Scharf et al., 2000; Pauly, 2000a). Yet, within-
species, the range of prey sizes consumed increases
with length, with maximum prey sizes increasing
while minimum prey sizes remain constant (Juanes
and Conover, 1994; Scharf et al., 2000). This is
the result of the fact that as fish grow their relative
success rates increase due to ontogenetic changes in
sensory and swimming capacities (e.g., Juanes and
Conover, 1994) as well as in mouth morphology,
and thus successively larger prey are included in the
diet. Yet smaller prey (i.e., of lower TROPH) is
never excluded, since this prey’s relative vulnera-
bility remains high (e.g., Juanes, 1994; Juanes and
Conover, 1994; Scharf et al., 2000). In fact, many
studies have demonstrated that fish tend to ingest
prey much smaller than the possible maximum: the
latter is estimated from their vertical mouth dimen-
sion (Schmitt and Holbrook, 1984; Hambright, 1991;
Juanes, 1994). One may postulate that this is also true
for among-species comparisons, as indicated by the
high variance in the plot of predator-prey length rela-
tionships for a large number of fish species (see Pauly,
2000a). The above-mentioned facts are in agreement
with an asymptotic relationship between body length
and TROPH for both among-species (Figure 8) and
within-species (Figure 10) comparisons.
There are three main methods for estimating
TROPH values, all of which display advantages and
disadvantages (Stergiou and Polunin, 2000): (a) from
stomach content data, with food items expressed in
terms of weight, volume or caloric content, with all
three measures being equivalent (Palomares and Sa-a,
1998); (b) from isotopic data (Kline and Pauly, 1998;
Pinnegar and Polunin, 1999, 2000; Vander Zanden
et al., 1999; Polunin and Pinnegar, 2000); and (c)
from existing empirical equations between TROPH
and body size (e.g., Pauly and Palomares, 2000) for a
wide range of fish species, from which TROPH can be
estimated from reported body sizes for data-poor situ-
ations. The latter is typical of the Mediterranean Sea
for which the regression equations shown in Figure
8 can be used. Finally, the Ecopath approach also
 247

Figure 10. Relationship between TROPH values and body length (midpoint of length class; in cm) for 11 fish stocks. (a) Trigla lyra in the
Western Mediterranean Sea, 1976–1978; (b) Chimaera monstrosa in the Western Mediterranean Sea, 1976–1978; (c) Caelorinchus caelorhincus
in the Western Mediterranean Sea, 1976–1978; (d) Merluccius merluccius in the Gulf of Lions, 1993; (e) Merluccius merluccius in the Central
Mediterranean Sea, 1982–1983 (the first length class was excluded from the analysis); (f) Pagellus acarne in the Evvoikos Gulf, 1987; (g)
Pagellus erythrinus in the Gulf of Lions, 1979–1980; (h) Pagellus erythrinus in the Central Tyrrhenian Sea, 1981–1982; (i) Lepidopus caudatus
in the Western Mediterranean Sea, 1976–1978; (j) Solea solea in the Western Mediterranean Sea, 1976–1978; and (k) Zeus faber in the Evvoikos
and Pagassitikos Gulfs, 1986–1988. The values of the parameters of the relationships are shown in Table 4.
248
provides an important tool for including additional
constraints on TROPH estimates derived from various
methods (Stergiou and Polunin, 2000).
The stomach content method, on which all the
studies reviewed here were based, assumes that all
major prey are identified and quantified correctly,
and their TROPH values are validated (Polunin and
Pinnegar, 2000; Stergiou and Polunin, 2000). In fact,
there are important problems with identifying many
types of organisms (e.g., gelatinous zooplankton and
detritus) (Polunin and Pinnegar, 2000). In addition,
carnivores may have empty stomachs most of the
time (e.g., Beyer, 1998), most stomach content data
often represent mere snapshots of diet, and prey’s
TROPHs are often not known (Pinnegar and Polunin,
1999, 2000; Polunin and Pinnegar, 2000, 2002).
Thus, cross-validation using different methods (e.g.,
Ecopath: Christensen and Pauly, 1992a, b; Pauly et
al., 2000b; 15 N:14 N ratios: Kline and Pauly, 1998;
Polunin and Pinnegar, 2000) is essential. For instance,
although the study of Kline and Pauly (1998) suggests
a good agreement between TROPH values estimated
from stomach content data and 15 N:14 N ratio data,
Badalamenti et al. (2000) note that analogous estima-
tions do not fully agree for Mullus barbatus in the Gulf
of Castellammare.
Finally, based on the sampling characteristics and
methodology used in the studies reviewed (Figures
2 and 3) and on the results of our analysis, we
conclude with some recommendations for future
stomach content studies: (a) Stomach content studies
must be based on monthly or seasonal samples, which
provide an overall pattern of feeding, as opposed
to studies based on circumstantial sampling (Figure
2b); (b) As pointed out earlier (Gonzales-Sanson
and Aguilar-Betancourt, 1983; Palomares and Sa-a,
1998), with the exception perhaps of fish larvae, which
consume food items that are all uniformly small,
frequency of occurrence, numerical percentage and
rank relative abundance, which are used extensively
(Figure 2c), are not good indicators of how much a
food item contributes to the diet of a given popula-
tion. Thus results should be reported in terms of either
volume or weight; (c) Researchers must always report
the length range together with the mean length (and
its SE) of the specimens studied (see Figure 2d); (d)
Researchers must report the total number of stomachs
analyzed and the percentage of empty stomachs, with
the former determined from the plot of number of prey
items vs. number of stomachs analyzed (e.g., Link
and Almeida, 2000) so that stomach content data is
representative of the full array of prey. In fact, the
analysis of a large food habit database, referring to
the shelf ecosystem of the NE United States, indi-
cates that for most species diet can be adequately
described based on 500–1,000 stomachs, with this
figure being generally smaller for food specialists and
larger for food generalists (Link and Almeida, 2000).
Such a range is much higher than those on which
the majority of the studies reviewed here were based
(Figure 3); (e) Researchers must estimate feeding
habits for different length classes (see Figure 10),
especially for consumers that are known to be fish
eaters; (f) Feeding studies should be accompanied by
estimates of TROPH values; and (g) The collection of
routine time series of stomach content data will allow
us to search for spatial and temporal trends in species’
TROPHs and for correlations with other parameters
such as fishing effort.
Acknowledgements
The authors wish to extend their gratitude to an
anonymous reviewer and Dr. D. Pauly for useful
suggestions and comments.
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