Journal of
Applied Ichthyology
J. Appl. Ichthyol. 28 (2012), 7581
 2011 Blackwell Verlag, Berlin
ISSN 01758659
Diet and feeding habits of Economidichthys pygmaeus (Perciformes: Gobiidae)
in Lake Pamvotis, NW Greece
By Ch. Gkenas1, S. Malavasi2 and I. Leonardos1
1
Department of Biological Applications and Technology, University of Ioannina, Ioannina, Greece; 2Department of Environmental
Sciences, Ca Foscari State University, Castello, Venezia, Italy
Summary
The purpose of the study was to dene the diet composition,
feeding preferences, seasonal, size and sex related changes in
diet, feeding strategy and diel cycle of Economidichthys
pygmaeus. Important features of the overall feeding patterns
of the goby include (i) feeding activity, (ii) bottom habits and
(iii) consumption of food items to identify the dietary breadth.
Stomach contents of 533 specimens, 13.4248.65 mm total
length (TL), collected by hand net (2 mm mesh size) from
October 2006 to September 2007 were analyzed. The percent-
age of 76 empty stomachs (14.26%) did not vary signicantly
with season. The food composition suggested that the goby is a
carnivorous sh. Prey items identied in the stomachs
belonged to four major groups: Crustaceans, Insects, Mollusca
and Plants. Copepods were the most important
(%IRI = 57.51), especially in sh larger than 23 mm TL.
Chironomids (%IRI = 43.23) constituted the main prey for
sh <23 mm TL. Diet composition showed little seasonal
variation. Copepoda dominated the diet in autumn and winter
(43.4%; 82.4%) and were replaced by Chironomidae larvae in
spring (39.7%) and summer (47.7%). During the reproductive
season, large males show a narrow food spectrum as opposed
to large females, reecting the dierent activity pattern
exhibited by males and females during the breeding season,
since the former supports parental care. According to the
modied Costello graphical method, specialist individuals
form the goby population that feeds on two preferential prey
types (e.g. Copepods, Chironomids). Nevertheless, they con-
sume some occasional prey (e.g. Cladocera, Insects, Gastro-
pods). The species foraged chiey at night and early morning,
during lower light intensity. The highest feeding activity
recorded at night (02.00 hours) and during early morning
(08.00 hours) could be related to the cryptic behaviour
displayed by the species.
Introduction
Among the Eastern Atlantic-Mediterranean gobioid shes, a
so-called sand goby group consisting of four phenetic genera
has been recognized: Pomatoschistus (Gill, 1863), Knipowits-
chia (Iljin, 1927), Gobiusculus (Duncker, 1928), and Econom-
idichthys (Bianco et al., 1987). The West Balkan genus
Economidichthys (Holly, 1929) comprises two endemic species:
E. pygmaeus and E. trichonis. Both species have a dermal
perianal organ, unique among teleosts, which may represent a
level of specialization higher than that of other teleosts
(Bianco et al., 1987; Economidis and Miller, 1990; Miller,
1990).
U.S. Copyright Clearance Centre Code Statement: 01758659/2012/28010075$15.00/0
Received: March 28, 2011
Accepted: August 25, 2011
doi: 10.1111/j.1439-0426.2011.01912.x
Economidichthys pygmaeus is an endemic small-bodied,
short-lived species restricted to northern and western Greece
(Miller, 2004). Specically, it is distributed in the Thyamis,
Louros, Arachthos and Achelloos rivers and in the Trichonis,
Lyssimachia, Ziros and Ozeros lakes. Recently it has been
recorded in Pamvotis Lake where it probably arrived from the
rivers Louros and or Kalamas (Thyamis) (Leonardos et al.,
2008). The species appears to be extinct on Lefkas Island
(Economidis, 1991) and is protected under Greek law (No.
67 1981). Greeces updated edition of the Red Book of
Endangered Species (2009) has evaluated its conservation
status, which is now assigned a least concerned species, i.e.
without facing any critical dangers (Economidis, 2009).
Economidichthys pygmaeus is generally considered a benthic
predator, feeding on benthic invertebrates and crustaceans
(Daoulas et al., 1993) but little information is available
regarding its diet. The aim of the present study was to provide
the rst information on the feeding ecology of E. pygmaeus.
Since sh feeding and prey availability vary temporally, the
sh diet needs to be described on a diel and seasonal basis
(Carman et al., 2006). More specically, the objectives
addressed are to: (i) provide information on the diet of
E. pygmaeus, with special emphasis on seasonal and ontoge-
netic dietary variation, as well as on its feeding strategy; (ii)
describe sex-related changes in the diet, particularly during the
breeding season; and (iii) examine seasonal diel variations in
the diet over a 24-h cycle.
Materials and methods
Study area
Lake Pamvotis, NW Greece (3940234N, 2053248E) is a
shallow Mediterranean lake with a mean depth of 4.5 m and
maximum depth of 7.5 m. The lake occupies an area of
22.8 km2 and is approximately 470 m above sea level. An
ancient lake ecosystem, Lake Pamvotis is situated in a
topographically diverse landscape on the western ank of the
Pindus mountain range (Tzedakis et al., 2002), and is of global
signicance in terms of biodiversity (Krystufek and Reed,
2004). The native sh fauna of the lake consists of only four
species (Pelasgus epiroticus, Luciobarbus albanicus, Squalius
pamvoticus and Anguilla Anguilla). Up until now, twenty exotic
or translocated sh species, representing eight families, have
been introduced into Lake Pamvotis. Among them, the pelagic
Rutilus panosi, benthic Silurus aristotelis, allogynogenetic
Carassius gibelio, and littoral E. pygmaeus and Gambusia
holbrooki have become established with viable and dense
populations (Leonardos et al., 2008).
76
Preliminary samplings show that the population of gobies is
currently in a restricted shallow area at the Perama site
(3941678N; 02051285E). This site is in a well-preserved a conversion of 0.27 mg of dry weight mm3 (Lindegaard,
area, 7 4 m, with maximum depth of 1.5 m. The littoral zone
is characterized by dense aquatic vegetation (Myriophyllum
spicatum) and the common reed (Phragmites australis). The
reeds provide shelter against predators (Gotceitas and Colgan,
1989; Savino and Stein, 1989) and act as goby spawning sites
(Daoulas et al., 1993). The substratum is mainly covered with
pebbles mixed with coarse sand. In addition to E. pygmaeus,
two other sh species occur in the sampling site: Knipowitschia
caucassica Berg, G. holbrooki Girard, and a shrimp species
Atyaephyra desmarestii Millet.
Sampling
Fish were collected monthly between October 2006 and
September 2007, using a hand net (2 mm mesh size) and
preserved in 4% formaldeyde solution for later laboratory
analysis. Sampling was carried out during the morning
(09.00 hours) along the littoral zone in the study site. Each
survey consisted of thirty hauls at the sampling station in a
total of 20 min.
In the same period, three water and sediment samples were
collected monthly in the littoral zone of the sampling site using
a P.V.C core of 130 cm height with a diameter of 10 cm to
analyze the food resources. The core penetrated 30 cm into the
sediment in a place where the water level reached about
100 cm. This manner ensured that sediment and water samples
maintained a constant volume (Caiola et al., 2001). The water
was then separated from the sediment and ltered through a
250 lm nylon sieve. A 500 ml aliquot of each sediment sample
was ltered through a 500 lm steel sieve. The invertebrates
retained in the net were collected and preserved in 4%
formaldehyde, sorted in the laboratory into taxonomic cate-
gories (order or family level) and counted. The percentage of
each taxonomic group was calculated based on the sum of all
invertebrates in the sample.
Feeding activity of E. pygmaeus was examined on a diel
cycle in a seasonal pattern. Gobies were collected in May,
August and October 2007 and January 2008. On each
sampling occasion, specimens were collected at 6-h intervals
over 24 h (08.00, 14.00, 20.00, 02.00 hours) with a hand net
(2 mm mesh size) at a depth of 0.51 m. Each collection lasted
about 30 min. Fish captured were immediately killed with an
overdose of MS-222, preserved in 4% formaldehyde and
transferred to the laboratory.
Laboratory methods
In the laboratory, specimens were measured to the nearest
0.1 mm using a digital caliper [total lengths (TL)]. All
individuals were eviscerated, sexed (generally from external
sexual dimorphism or by macroscopic examination of the
gonads), and weighed (total and eviscerated weight to the
nearest 0.1 mg using a digital balance). Contents of each
stomach were washed in a petri dish, where prey items were
identied, separated and counted. Measurements of length,
width and depth were taken for each individual prey item using
a dissecting stereoscope with an ocular micrometer. The
volume of food items was estimated by calculation of the
volume of a geometric solid of similar size and shape (Bowen,
1983). The volume of uncountable categories (plant leaves) in
the gut contents was estimated to the nearest 0.00025 mm3
Ch. Gkenas, S. Malavasi and I. Leonardos
with a Neubauer counting chamber (Garcia-Berthou, 1999).
Estimated wet volume was later transformed to biomass using
1992; Alcaraz and Garcia-Berthou, 2007). Fragments of
organisms, such as the heads, were considered to be individ-
uals. Prey composition was identied to the lowest possible
systematic level.
Data analysis
Prey items were identied and classied; those food items
representing <5% in frequency of occurrence were pooled
into families or orders. The seasonal feeding activity pattern
was derived from the seasonal variation of percentage of
empty guts following Caiola et al. (2001). The contribution of
each prey (dietary item) to the diet was estimated using three
simple indices [number percentage (%N), volume percentage
(%V) and frequency of occurrence percentage (%FO)] dis-
cussed and revised by Hyslop (1980) and Cortes (1997). The
index of relative importance (IRI) (Pinkas et al., 1971)
standardized to %IRI (Cortes, 1997), which combines the
values of the simple indices mentioned above, was also used as
the best approach to gain an adequate description of the diet
(Hyslop, 1980) and allowed comparison between dietary items
of the same species and between diets of dierent species
(Cortes, 1997).
In order to analyze seasonal and intraspecic dierences in
diet, the months were categorized into seasons (autumn =
SeptNov; winter = DecFeb; spring = MarMay; sum-
mer = JunAug) and species were divided into three size-
groups: small individuals (TL <23 mm), intermediate males
(IntM) females (IntF) (TL = 2330 mm) and large males
(LM) females (LF) (TL > 30 mm).
The diet overlap among seasons and size groups was
estimated
P using the Schoener (1970) similarity index: a = 1
0.5 |pijpik|, where pij is the proportion of resource
i consumed by group j (e.g. males) and pik is the proportion
of resource i consumed by group k (e.g. females). Volumetric
(V) diet measurements were used in the calculation of this
index. Zero value indicates no overlap, while 1 represents
complete overlap. The index value is generally considered
biologically signicant when it exceeds 0.6 (Wallace, 1981).
Prey diversity in the diet, which is a measure of trophic niche
breadth (Scrimgeour and Winterbourn, 1987), P was calculated
using the ShannonWeaver index (H): H = ) pi lnpi, where
pi is the proportion of individuals of prey type i (Shannon and
Weaver, 1963). This index has adequate sensitivity for detect-
ing changes in species diversity and provides a general
indication of the relative magnitude of trophic specialization
(Berg, 1979).
Following Lechowicz (1982), comparison between diet
composition and resource availability was made using the
Vanderploeg and Scavia (1979) relativized electivity index
(E*):
X
Ei Wi  n1 Wi n1 1 ; whereWi ri pi1 ri pi1 1 ;
ri is the relative (proportional) abundance of prey i in the diet,
pi is the relative abundance of prey i in the environment and n
is the number of prey types included in the analysis. This index
ranges from +1 (positive selection) to )1 (negative selection);
values near zero indicate neutral selectivity. Although other
electivity indices such as those of Ivlev or Strauss are in wider
use, they have several undesirable properties and that of
Diet of Economidichthys pygmaeus 77

Vanderploeg and Scavia (1979) is the single best electivity Table 1

index (Lechowicz, 1982). Diet composition of 533 E. pygmaeus (%FO, percentage frequency of
occurrence; %N, percentage numerical abundance; %M, percentage
To determine the feeding strategy along the annual cycle volumetric composition; % IRI, index of relative importance on a
studied, the modied Costello graphical method (Amundsen percent basis (%IRI)
et al., 1996) was used where the prey-specic abundance (Pi),
dened as the percentage a prey taxon comprises in only those Prey category %FO %N %V %IRI
predators in which the actual prey occurs, is plotted against the
Crustacea
frequency of occurrence (Fi) on a two-dimensional graph. Copepoda 64.8 50.3 19.7 57.4
Information on prey importance and feeding strategy of the Daphnia sp. 12.5 6.6 9.4 2.5
predator can be obtained by examining the distribution of Bosmina longirostris 2.0 0.5 0.1 0.1
points along the diagonals and the axes of the diagram Alona rectangula 23.9 8.0 0.8 2.7
Chydorus sphaericus 4.6 0.9 0.2 0.1
(Amundsen et al., 1996; Caiola et al., 2001). Ostracoda 9.0 2.4 0.8 0.4
Diel feeding activity was assessed by comparing the volume Insects
data of prey categories for all gobies. Because the ANOVA Chironomidae larvae 56.7 17.9 22.4 29.0
assumptions of normality and homogeneity of variance were Diptera pupae 6.2 1.1 13.7 2.7
not met, seasonal and diel dierences in the volume of stomach Ephemeroptera larvae 3.4 0.5 6.8 0.7
Hymenoptera larvae 1.7 0.1 2.1 0.1
contents were tested with the non-parametric KruskalWallis Other insects (n.i) 2.1 0.2 1.4 0.1
test. MannWhitney U-tests with a Bonferroni adjustment Mollusca
were used to examine all pair-wise comparisons for signicant Sphaeriidae 2.4 0.5 6.2 0.2
dierences. A chi-square test was also applied to determine Planorbidae 3.2 0.7 15.4 0.7
Plant materials 26.7 8.9 0.2 3.0
whether the percentage of empty guts was signicantly
Unidentied 9.6 1.6 0.8 0.3
dierent between time intervals. All data analyses were
performed with SPSS 19 for Mac OS. n.i., non-identied material.

Copepoda dominated the diet in autumn and winter (43.4

Results
Seasonal variation in feeding activity
The seasonal variation in the percentage of empty alimentary
tracks did not reect any marked discontinuity in E. pygmaeus
feeding activity. In Lake Pamvotis, the seasonal percentages of
empty alimentary tracks were 10.3% in autumn, 10% in
winter, 15.7% in spring and 20.2% in summer (G-test = 7.71,
d.f. = 3, P = 0.052).
Overall diet composition
In total, 533 E. pygmaeus stomachs were examined of which
14.26% (n = 76) were empty. Analysis of stomach contents
identied 14 prey categories belonging to four major groups:
Plants, Crustaceans (Copepods, Cladocerans, Ostracoda),
Insects (Chrironomidae larvae, Diptera pupae, Ephemeroptera
larvae, Hymenoptera larvae), and Mollusca represented
by Gastropoda (Planorbidae) and Bivalvia (Sphaeriidae)
(Table 1). Copepoda were the principal component of the diet
present in 64.8% of the sh, comprising 50.3% of the diet by
number, 19.7% by volume and 57.4% by IRI. Chironomidae
larvae were the second most common prey item (56.7% F) and
accounted for 17.9% by number, 22.4% by volume and 29%
by IRI. When percentage by volume (V) was considered,
Insects (Chironomidae larvae 22.4%; Diptera pupae 13.7%;
Ephemeropteran larvae 6.8%) were the most important prey
categories, followed by Copepoda 19.7%, and Mollusca
(Planorbidae 15.4%; Sphaeriidae 6.2%). Some littoral
Cladocerans, particularly Daphnia sp. and Alona rectangula
were slightly more common (12.5% F; 23.9% F) but less
important in terms of number (6.6%; 8%), volume (9.4%;
0.8%) and IRI (2.5%; 2.7%).
Seasonal variation in diet composition and feeding pattern
The importance of major prey in the four seasons was similar
to the pattern in the total sample (Table 1). The IRI (%) of the
dierent food items throughout the year are shown in Table 2.
82.4%, respectively) and were replaced by Chironomidae
larvae in spring (39.7%) and summer (47.7%). In spring and
summer, there was an increase in the importance of Cladocera
(24.8 and 19.8%, respectively). Apart from those mentioned in
the seasonal diet analysis, aquatic insects increased in impor-
tance in spring (8.9%) and summer (3.9%). Plant materials
and Mollusca were sporadically important in autumn (7.7%)
and spring (3.9%).
Gobies showed a narrow diet breadth for prey utilization as
a result of their specialization in Copepods and Chironomids
(Table 3). In winter, a higher number of prey items was
observed, primarily due to the dominance of Copepods. The
spring samples had the highest prey diversity values
(H = 0.56) while the summer samples had the lowest
(H = 0.44). No prey item was absent during the year cycle.
Niche overlap results in Table 4 showed that E. pygmaeus did
not consume the same prey type in all seasons. The lowest
value was observed between autumnspring (a = 0.47) and
the highest value between autumnsummer (a = 0.76).
Intraspecic variation in diet composition and feeding pattern
The diet of small gobies was based on Chironomidae larvae
(43.2%), followed by Cladocera (29.1%), Copepoda (22.5%)
and, of lesser importance, aquatic insects (4.4%). The inter-
mediate males fed mainly on Copepods (60.9%), followed in
importance by Chironomids (22.8%), Cladocera (5.5%) and
aquatic insects (5.1%). The diet of intermediate females was
very similar to that of the preceding ones. In both large male
and large female individuals, Copepoda was the dominant and
most consumed prey (55.8 and 50.6%, respectively). Chiro-
nomidae larvae consumption was more important in large
females (26.5%) than in large males (17.9%). Cladocera
consumption was similar in both groups. Moreover, the
importance of aquatic insects and Mollusca appeared rela-
tively higher in relation to small size-classes (Table 2). The
importance of plant materials in the examined stomach
contents increased considerably in large males (8.8%).
78 Ch. Gkenas, S. Malavasi and I. Leonardos

Table 2
Seasonal and Intraspecic diet composition of E. pygmaeus based on %IRI values of major prey groups (n, number of digestive tracks;
Chironomidae (l.), Chironomidae larvae; IntM IntF, Intermediate males Intermediate females; LM LF, Large males Large females)

Season Size class of sh

Small IntM IntF LM LF

Autumn Winter Spring Summer (<23 mm) (2330 mm) (2330 mm) (>30 mm) (>30 mm)
Prey category n = 145 n = 131 n = 134 n = 123 n = 103 n = 113 n = 134 n = 82 n = 101

Copepoda 43.4 82.4 20.6 25.3 22.5 60.9 55.6 55.8 50.6
Cladocera 4.6 4.6 24.8 19.8 29.1 5.5 6.7 8.3 11.5
Chironomidae (l.) 42.6 8.1 39.7 47.7 43.2 22.8 34.5 17.9 26.5
Aquatic Insects 0.9 1.3 8.9 3.9 4.4 5.0 0.1 6.5 4.8
Ostracoda 0.1 0.3 1.2 0.2 0.2 0.5 0.6 0.1 0.6
Mollusca 0.2 1.6 3.9 0.5 0.1 1.1 0.1 2.4 4.4
Plants 7.7 1.7 0.7 1.9 0.3 3.6 2.2 8.8 1.5
Unidentied 0.6 0.1 0.3 0.7 0.2 0.5 0.3 0.2 0.2

Table 3 benthic prey such as Gastropoda (Planorbidae). Intermediate

Dietary breadth of E. pygmaeus expressed in terms of the Shannon females had dierent diets compared to intermediate males
Wiener index (H) and number of prey items in stomachs of specimens,
classied according to sampling season and size group (n, number of (a = 0.51), large males (a = 0.48) and large females
digestive tracks; IntM IntF, Intermediate males Intermediate fe- (a = 0.48). The frequent utilization of bigger prey items (e.g.
males; LM LF, Large males Large females) Daphniidae), hard-bodied insects and avoidance of very small
littoral crustaceans (A. rectangula, Chydorus sphaericus) by
Number of intermediate males and large gobies accounted for the dier-
Season n prey items H
ence. Food niche overlap was slightly lower between interme-
Autumn 145 785 0.52 diate and large females (a = 0.58).
Winter 131 1249 0.45
Spring 134 671 0.56
Summer 123 491 0.44 Prey selection
Size class of sh
Small (<23 mm) 103 421 0.43 To study prey preferences of E. pygmaeus, E* value was
IntM (2330 mm) 113 509 0.49 calculated to assess the attraction (positive values), the
LM (>30 mm) 82 719 0.53 avoidance, or the inaccessibility of prey (negative values) in
IntF (2330 mm) 134 764 0.49
comparison to prey abundance in the eld (Fig. 1). The high
LF (>30 mm) 101 783 0.57
E* values of Copepoda (0.95) and Chironomids (0.88)
indicated a strong preferential prey selection. Positive E*
Animal diet breadth for juveniles and intermediate size- values were found for C. sphaericus (0.64), A. rectangula
groups tended to be narrower than in the larger size groups (0.47), Planorbidae (0.47) and Diptera pupae (0.4). In contrast,
(Table 3). Large males (H = 0.53) and large females a negative E* was found for B. longirostris ()0.53) and
(H = 0.57) showed high diversity index values. The dietary Sphaeriidae ()0.36), indicating avoidance. Furthermore,
overlap varied consistently with the dierences in food choice gobies showed a neutral selection for Daphnia sp. (0.15),
recorded between length classes (Table 4). A signicant niche Plants (0.1), and Hymenoptera larvae (0.05).
overlap was noted between juveniles and intermediate males
(a = 0.66) and with intermediate females (a = 0.66), respec-
Feeding strategy
tively. The diet of large males and large females showed highly
signicant overlap (a = 0.9), consisting mostly of Copepods The relationship between prey specic abundance and fre-
and Chironomidae larvae and to a lesser extent of other quency of occurrence suggested some feeding variation. The
pelagic zooplankton dominated by Cladocera and littoral diagram of the modied Costello graphical method revealed

Table 4
Food overlap coecients (Schoeners index) of diet between seasons and size classes, E. pygmaeus (n, number of digestive tracks; IntM IntF,
Intermediate males Intermediate females; LM LF, Large males Large females)

Season n Autumn Winter Spring Summer

Autumn 145 0.57 0.47 0.76*

Winter 131 0.65* 0.49
Spring 134 0.52
Summer 123
Size class of sh Small (<23 mm) IntM (2330 mm) LM (>30 mm) IntF (2330 mm) LF (>30 mm)

Small (<23 mm) 103 0.66* 0.58 0.66* 0.59

IntM (2330 mm) 113 0.69* 0.51 0.74*
LM (>30 mm) 82 0.48 0.9*
IntF (2330 mm) 134 0.48
LF (>30 mm) 101

*signicant similarity for values of C 0.60

Diet of Economidichthys pygmaeus 79

1 20.00 hours (30%) than in the other time intervals

0.8 (08.00 = 14%, 14.00 = 23%, 02.00 = 12%) (v2 = 13.81,
d.f. = 3, P < 0.001). KruskalWallis tests indicated that total
0.6
volume varied signicantly between seasons (KruskalWallis
0.4 ANOVA, n = 317, P < 0.001) and throughout the day (Krus-
0.2 kalWallis ANOVA, n = 317, P < 0.001) (all data pooled).
0 MannWhitney pairwise comparisons, with a Bonferroni
adjustment for an overall condence level of 0.05, indicated
0.2
that the prey volume in goby stomachs was greater at night
0.4 than during the day (Fig. 3). Variations always occurred
0.6 between the intervals, with early morning samples having
0.8
signicantly higher values than those during daytime (Mann
Whitney U-test, n = 154, P < 0.05) and evening (Mann
1
Whitney U-test, n = 160, P < 0.05). Prey volume in night
Plant materials

Daphnia sp.

B. longirostris

A. rectangula

C. sphaericus

Copepoda

Ostracoda

Chironom. (l.)

Diptera (p.)

Ephem. (l.)

Hymen. (1.)

Sphaeriidae

Planorbidae
samples was signicantly heavier than those in the daytime
(MannWhitney U-test, n = 157, P < 0.05) and evening
(MannWhitney U-test, n = 163, P < 0.05). Early morn-
ing night samples (MannWhitney U-test, n = 163,
Fig. 1. Vanderploegs prey selectivity for various food components P > 0.05) and daytime evening samples had similar prey
found in E. pygmaeus guts. Positive values = prey selection, negative volume values (MannWhitney U-test, n = 154, P > 0.05).
values = prey avoidance. Non-selective feeding = zero values. Chi-
ronom. (l.), Chironomidae larvae; Diptera (p.), Diptera pupae; Ephem.
(l.), Ephemeroptera larvae; Hymen. (l.), Hymenoptera larvae
Discussion
two groups (Fig. 2). The rst group was represented mostly by The feeding pattern of the E. pygmaeus population is charac-
Copepoda followed by Chironomidae larvae, indicating that terized by consumption of Copepods associated with Clado-
there is some specialization toward one prey (Copepoda), as it cera, insect larvae and pupae, Mollusca and Plant materials.
is the only prey point positioned in the upper right part of the Copepoda was the most common prey in terms of abundance
graph. Copepods showed a higher abundance and occurrence and occurrence. However, aquatic insects (mostly Diptera
than the other prey items, being consumed by more than 60% pupae) and Chironomidae larvae were the dominant prey in
of shes, and constituting a dominant prey item. The second terms of weight. Benthic prey items found in the diet showed a
group found in the lower left part constituted most of the prey tendency of gobies to predate at the bottom, but not always,
items (e.g. A. rectangula, Daphnia sp. Ostracoda, Planorbidae,
Diptera pupae), indicating that these are rare or less frequent
prey (low frequency and abundance) and only consumed (a)
occasionally (generalistic feeding strategy).

Diel feeding pattern

The chi-square test revealed signicant dierences in the
percentage of empty guts between the time intervals. The
percentage of empty guts was signicantly higher at

70
Copep.

60

50

40
(b)
Pi(%)

Chironom. l.

Daph. sp. A. rectangula

30
Ostrac. Plants
Planorb.
20
B. longirostris
C. sphaericus
Sphaer.
10 Diptera p.
Ephem. l.
O. insects
Hymen. l.
0
0 10 20 30 40 50 60 70
Fi(%)

Fig. 2. Relationship among prey specic abundance (%Pi) and

frequency of occurrence (%FO) of food categories in E. pygmaeus
diet. Plots based on modied Costello graphical method (Amundsen
et al., 1996). (Copep., Copepoda, Chironom. l., Chironomidae larvae,
Daph. sp., Daphnia species, A. rectangula, Alona rectangula; Ostrac.,
Ostracoda; Planorb., Planorbidae; B. Longirostris, Bosmina longiros-
tris; Sphaer., Sphaeriidae; C. Sphaericus, Chydorus sphaericus; Diptera
p., Diptera pupae; Ephem. l., Ephemeroptera larvae; Hymen. l., Fig. 3. Mean total volume of food in goby stomachs. (a) Seasonal
Hymenoptera larvae; O. insects, Other insects) variation, (b) Diel variation at 6-h intervals. Error bars, SE
80
because one of the major prey (Copepoda) was found in the
nectonic organisms. Hence, it appears that E. pygmaeus was
not feeding exclusively on benthic prey but also on organisms
from the water column. Although the prey composition of
P. microps (Salgado et al., 2004; Leitao et al., 2006), P. min-
utus (Hamerlynck and Cattrijsse, 1994), Knipowitschia cauca-
sica (Kevrekidis et al., 1990) and P. bonelli (Bo et al., 2010)
includes polychaete radioles, mysids, small shrimps, amphi-
pods and aquatic insects apart from small crustaceans and
bivalves, E. pygmaeus has been reported to feed predominantly
on crustaceans and benthic invertebrates (Daoulas et al.,
1993), which was conrmed by the results of this study.
In all seasons, the proportion of empty stomachs showed a
lack of signicant dierences in feeding activity. The fact that
gobies maintained the same feeding activity in Lake Pamvotis
may be due to the stable climatic conditions throughout the
year (Kagalou et al., 2006; Ch. Gkenas, personal observa-
tions). Thus, gobies were able to exploit the available food
resources in order to satisfy their annual metabolic needs. Sand
gobies are considered opportunistic carnivores, feeding on
organisms selected on the basis of their availability (Pihl,
1985). Hence, the changes in the seasonal variation observed in
the consumption of some food types by E. pygmaeus may
suggest that diet reected prey availability. Examples are the
consumption of Chironomids in summer and higher abun-
dance of Copepods observed in the winter diet of the goby,
when densities of this prey type are higher (Kagalou et al.,
2006; Antonopoulos et al., 2008). The observed greater dietary
diversity in spring could be attributed to the possible decrease
in availability of particular prey during this season, which
would have forced E. pygmaeus to forage on other food types
(Chironomids, Cladocerans, aquatic insects) that might have
been readily available. Furthermore, the prey food diversity
was at its highest in spring, probably in an eort to
compensate the increase energy demands of the reproductive
period.
The ontogenetic switch in feeding habits is a general
phenomenon among sh as a result of increasing body size.
This occurs due to a shift from a diet of mostly small soft-
shelled prey, such as microcrustaceans, Chironomidae larvae,
to a more diverse adult diet which also includes larger prey,
such as Gastropods (Planorbidae), large Cladocerans and
aquatic insects (mostly Diptera pupae and Ephemeroptera
larvae). The wider dietary diversity in larger gobies, as
expressed by the ShannonWiener index, could reect the
ability of large specimens to utilize a wider range of habitat
resources, both in the trophic dimension (diet composition),
due to their increased morphological adaptations, as well as in
the spatial dimension (distribution), due to their greater
mobility and hence dispersal rate (Vassilopoulou, 2006). Large
males exhibited a narrower food niche than large females. This
may be due to the fact that males were probably limited to the
food available around the nest since they display territorial
behaviour and provide parental care for the eggs in the
breeding season (Daoulas et al., 1993; Gkenas et al., 2010).
Females apparently behave as food maximisers, feeding on a
high number of benthic prey categories such as Chironomids,
Gastropods and Cladocerans. A similar change in the energy
requirements for reproduction and growth was suggested for
P. minutus by Salgado et al. (2004), with gobies showing a
preference for Polychaetes, Isopods and some Amphipods
compared to Copepods because of their richer caloric
content. No goby eggs were observed in the gut contents,
which according to Hamerlynck and Cattrijsse (1994) may
Ch. Gkenas, S. Malavasi and I. Leonardos
suggest that nest sites were not a limiting factor in this lake. In
spite of diet and prey breadth variations, small to intermediate
size gobies and large males and females exploited essentially
the same prey base, showing consistently high overlapping. On
the other hand, the low diet overlap between intermediate
females intermediate males to adult gobies could represent an
intraspecic segregation in the dierential consumption of the
same prey items due to the dierent dimensions reached by
medium size female gobies.
Electivity depended on prey availability and sh size.
Overall, smaller sh selected smaller prey such as littoral
Cladocerans (C. sphaericus and A. rectangula), while larger
sh showed a greater preference for diverse larger prey
(Insects, Gastropods). Such a diet may be protable in terms
of trade-os between item size, body hardness and ease of
catching. The high E values for some food categories, such as
Copepods and Chironomidae larvae, suggested the importance
of these food items as a food resource. The negative E values
for Bivalvia (Sphaeriidae) could be explained by the fact that
gobies with no specialized jaws usually avoid Bivalves (Zander,
1979).
On analysing the diet composition, one can easily conclude
that E. pygmaeus has a homogeneous diet strategy and a
narrow niche width. The modied Costello method revealed a
focus on two preferential prey types following a generalist
feeding strategy with an adaptation to seasonal prey avail-
ability. At the study area, this predator specialized in Cope-
pods and Chironomids, with other prey taken only
occasionally.
The results revealed a clear distinction between seasonal
changes of feeding activity, since diet shifts of E. pygmaeus
can be interpreted as a reaction to changing food availability.
Also, stomach contents of gobies indicated a distinct 24-h
rhythm of feeding. E. pygmaeus appeared to forage chiey at
night and early morning at times of lower light intensity. The
latter conrmed the lowest empty stomachs of gobies during
the night and early morning hours. This feeding behaviour
could be a fright response to sudden switches between light
and darkness because gobies display a cryptic behaviour
burrowing into sediment and exhibiting sit and wait hunting
(Miller, 2004). In similar diel studies, P. microps and P. min-
utus were more active during the day, with peaks in activity
occurring mainly at dawn to sunrise, and to a lesser extent in
the afternoon (del Norte-Campos and Temming, 1994).
In conclusion, this paper contributes a rst insight into the
diet and feeding strategy of E. pygmaeus, whose diet in Lake
Pamvotis consists of various benthic macroinvertebrates and
crustaceans. E. pygmaeus has a specialized zooplanktivorous
diet (especially Copepods) in all size classes and seasons. This
species feeds chiey at night and early morning, during the
lower light intensity. Feeding intensity along the diurnal cycle
probably corresponds to the cryptic behaviour displayed by
the species.
Acknowledgement
The authors are grateful to Callie Kalogerson for the linguistic
improvement of the manuscript.
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Authors address: Ioannis Leonardos, Department of Biological
Applications and Technology, University of Ioann-
ina, Ioannina, PC, GR-45 110, Greece.
E-mail: ileonard@uoi.gr