Professional Documents
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1
Kagawa Prefectural Fisheries Experimental Station, Takamatsu, Kagawa 761-0111 and 2Faculty of
Biotechnology, Fukui Prefectural University, Obama, Fukui 917-0003, Japan
ABSTRACT: Feeding habits of Favonigobius gymnauchen, Repomucenus spp. and Tarphops oli-
golepis were examined during the period from May to August in 1999, 2000 and 2001 at a sandy
beach in the central Seto Inland Sea, Japan. The three species of fish accounted for more than 60%
of all fish collected in terms of number of individuals. All three species mainly consumed small crus-
taceans. However, the major prey of F. gymnauchen and Repomucenus spp. differed from that of
T. oligolepis. F. gymnauchen and Repomucenus spp. mainly consumed mysids and small crangonid
shrimp (<12 mm in body length), which predominated in the study area. T. oligolepis actively selected
only epifaunal mysids Nipponomysis ornata and avoided crangonid shrimp and gammarids. The
frequencies of occurrence of fish in the guts of the three dominant species were very low, and larval
and juvenile Japanese flounder were not observed in any of the three fish species. The diet of juvenile
Japanese flounder was similar to that of the three species. From these results, it appears that these
three fish are competing species for the flounder.
KEY WORDS: demersal fish, Favonigobius gymnauchen, feeding habits, food web, Repo-
mucenus spp., sandy beach, Seto Inland Sea, Tarphops oligolepis.
measured for each day with Pianka’s O index,35 Of which, F. gymnauchen, Repomucenus spp.,
which was as follows: P. olivaceus and T. oligolepis comprised 64.7% of
the total number and 95.6% of the total weight
O12 = Σp1ip2iΣ(p1i2p2i2)(1/2),
of fish collected. Because sizes of R. ercodes and
where p1i and p2i are the proportions of the number H. japonica were small (major size: <15 mm in
of prey item i in predator 1 and predator 2, respec- TL), the stomachs of two species were not
tively. This index ranges from 0 (no overlap) to 1 examined. Although large dragonets (larger
(complete overlap). Here, we used the stomach than approximately 40 mm in TL) were composed
contents of the Japanese flounder (9.80–75.95 mm of R. richardsonii and R. beniteguri (mostly
TL) collected with the three dominant species.23 R. beniteguri), smaller individuals were not identi-
fied. Developmental stages of all of specimens
of F. gymnauchen, Repomucenus spp. and
RESULTS T. oligolepis were juveniles or adults.
Besides fish, 32,235 mysids (wet weight: 166.1 g)
Abundance of fish, mysids and crangonid shrimp and 120,491 crangonid shrimp Crangon uritai
(3591.9 g) were also collected. The dominant
A total of 4460 individual fish, including more than mysids were Iiella ohshimai and Nipponomysis
25 species in 15 families, were collected by 35 ornata.
operations of the 2-meter beam trawl at Ohama
Beach from May to August in 1999, 2000 and
2001 (Table 1). In terms of individual numbers, Gut contents
F. gymnauchen (percentage in number: 50.7%),
Rudarius ercodes (26.1%), Repomucenus spp. Of 150 individuals of F. gymnauchen examined,
(7.0%), Heteromycteris japonica (4.7%), P. olivaceus which ranged from 20.7 to 71.4 mm in TL, 76.7%
(4.3%) and T. oligolepis (2.7%) were dominant. contained food (Table 2). In Repomucenus spp.,
Table 1 Demersal fish collected by a 2-meter beam trawl at Ohama Beach from May to August in 1999, 2000 and
2001
Family Species Number of individuals Weight (g)
Clupeidae Konosirus punctatus 3 0.1
Syngnathidae Syngathus schlegeli 3 0.2
Hippocampus mohnikei 3 0.4
Scorpaenidae Sebastes inermis 1 1.8
Inimicus japonicus 2 6.8
Congiopodidae Hypodytes rubripinnis 18 3.9
Platycephalidae Platycephalidae spp. 80 3.2
Cottidae Pseudoblennius cottoides 2 1.6
Sparidae Pagrus major 7 0.5
Acanthopagrus schlegeli 31 2.2
Labridae Halichoeres poecilopterus 3 94.2
Blenniidae Scartella emarginata 2 0.1
Callionymidae Repomucenus spp. 313 214.7
Gobiidae Sagamia genetonema 5 2.0
Favonigobius gymnauchen 2262 5612.7
Tridentiger trigonocephalus 2 0.1
Gobiidae spp. 24 0.4
Paralichthyidae Paralichthys olivaceus 190 73.2
Tarphops oligolepis 122 72.4
Soleidae Heteromycteris japonica 211 122.0
Pseudaesopia japonica 1 –
Monacanthidae Rudarius ercodes 1166 33.9
Thamnaconus modestus 1 0.6
Tetraodontidae Takifugu niphobles 2 0.1
Unidentified 6 0.7
Total 4460 6247.9
84.4% of 90 individuals examined contained food. of T. oligolepis at the same total length of larger
Although this species (n = 313) ranged from 20.1 to than 15.0 mm.
107.4 mm in TL, few individuals ranging from 30 to
40 mm were present (n = 5). The feeding incidence
(percentage of stomach with food) of 122 individ- Diet shift with growth
uals of T. oligolepis ranging from 9.0 to 75.4 mm TL
was 87.7%. The feeding incidences of these three The reverse trends of diet shift with growth were
species were not significantly correlated with body observed in mysids and crangonid shrimp in
size (Spearman’s rank correlation coefficient test, F. gymnauchen (Fig. 1). That is, the weight com-
P > 0.3). position of mysids decreased with total length,
The main prey items in the three dominant whereas that of crangonid shrimp increased. The
fish, F. gymnauchen, Repomucenus spp., and diet of small Repomucenus spp. was composed
T. oligolepis, were small crustaceans (mostly of mysids and gammarids; and with growth,
mysids, gammarids and decapods) (Table 2). Set- the percentage of crangonid shrimp and fish
tled larval and juvenile Japanese flounder were increased. The stomach contents of T. oligolepis
not observed in the guts of any fish examined. consisted primarily of mysids of all sizes, and no
F. gymnauchen mainly consumed mysids (%IRI: ontogenetic shift in the feeding habits was
67.9%) and crangonid shrimp (26.0%). The %IRI observed.
of I. ohshimai (30.5%) was higher than that of The size relationships among the three fish and
N. ornata (20.4%). Other prey items were gam- their major prey items are shown in Figure 2. In
marids, brachyurans, fish and eggs of Hemiram- F. gymnauchen up to 40 mm, Repomucenus spp.
phus sajori, but the %IRI of these prey items were up to 50 mm and T. oligolepis up to 30 mm, the
very low. In the gut of Repomucenus spp., mysids body length (BL) of mysids consumed increased
and crangonid shrimp predominated, and sand with the total length of the predators. The maxi-
grains were often observed (frequency of occur- mum prey size of F. gymnauchen, Repomucenus
rence: approximately 40%). T. oligolepis mainly spp. and T. oligolepis were 9.3 mm (mean: 5.9 mm),
consumed mysids, particularly N. ornata. 10.0 mm (6.4 mm) and 8.3 mm (4.7 mm), respec-
The dietary diversity (Simpson diversity index, tively. The maximum size of mysids (approximately
1 − D) values of F. gymnauchen and Repomucenus 9 mm BL) was consistent with that of mysids in the
spp., of 0.81 and 0.80, respectively, were higher environment.23 The maximum size of crangonid
than that of T. oligolepis, 0.66. The dietary diversity shrimp increased with the growth of the predators,
of Japanese flounder P olivaceus calculated (not and the mean body lengths of shrimp observed in
tabled) was 0.77. the guts of F. gymnauchen, Repomucenus spp. and
T. oligolepis were 8.3 mm (range: 5.2–11.8 mm),
8.5 mm (5.0–11.3 mm) and 7.7 mm (3.8–10.2 mm),
Relationship between mouth size and fish length respectively. In gammarids, it was not possible to
analyze the size relationship between predator and
Mouth size (upper jaw length and/or mouth width) their prey because of lack of prey body length.
was significantly correlated with total length However, the mean body length of the gammarids
(Table 3). The slopes of the regression lines were eaten by F. gymnauchen, Repomucenus spp. and
significantly different between P. olivaceus and T. oligolepis were 2.3 mm (range: 0.8–3.8 mm),
T. oligolepis (analysis of covariance, P < 0.01), and 1.3 mm (0.8–1.8 mm) and 2.2 mm (0.8–4.2 mm),
the mouth size of P. olivaceus was larger than that respectively.
Table 3 Relationships between upper jaw length (UJL: in mm)/mouth width (MW: in mm) and total length (TL: in mm)
of Favonigobius gymnauchen, Repomucenus spp., Tarphops oligolepis and Paralichthys olivaceus. Regression equations
and significance levels were shown
Equation Degrees of freedom r2 F P
Favonigobius gymnauchen UJL = 0.0523 TL + 0.209 49 0.81 207 <0.01
MW = 0.0913 TL − 0.00068 49 0.83 251 <0.01
Repomucenus spp. MW = 0.0646 TL + 0.149 49 0.94 766 <0.01
Tarphops oligolepis UJL = 0.0678 TL + 0.600 29 0.90 273 <0.01
Paralichthys olivaceus UJL = 0.103 TL − 0.0704 29 0.97 1034 <0.01
Feeding ecology of dominant fish at a beach FISHERIES SCIENCE 1337
80
60 All of the mean overlap values, Pianka’s O index, for
pair species from F. gymnauchen, Repomucenus
40
spp. T. oligolepis and P. olivaceus ranged from
20 0.48 to 0.66, although the individual values greatly
0 fluctuated (Fig. 3). Those values for pairs of
0 20 30 40 50 60 70 76 F. gymnauchen and Repomucenus spp., T. oligolepis
Tarphops oligolepis and P. olivaceus were relatively higher.
42 25 17 13 4 4 2
100
Composition (%)
80 DISCUSSION
60
Gut contents and prey selection
40
20 The feeding incidences of the three dominant
0 fish, F. gymnauchen, Repomucenus spp. and
0 20 30 40 50 60 70 110 T. oligolepis, varied between 76.7 and 87.7% during
Total length (mm) the day. The major prey items were the epifaunal
Mysidae Callianassdae mysids N. ornata and A. ijimai, and the sand-
burrowing mysid I. ohshimai. I. ohshimai, Archae-
Gammaridea Pisces omysis kokuboi and A. japonica, belonging to the
subfamily Gastrosaccinae, burrow under the sand
Crangonidea Others during the day and swim up to the water column
during the night.36,37 The stomach content index
Fig. 1 Wet weight composition of stomach contents (SCI) of many species of juveniles in the surf-zone
by size for Favonigobius gymnauchen, Repomucenus of the Pacific Ocean, which feed on the sand-
spp. and Tarphops oligolepis. Numerals above bars burrowing mysids A. kokuboi, was increased in
indicate the number of specimens examined. the night.22 Imada and Nanba24 observed that
Body length of prey (mm)
Table 4 Selectivity (Chesson’s αi) of the major food items for Favonigobius gymnauchen, Repomucenus spp. and Tar-
phops oligolepis at Ohama Beach in 1999, 2000 and 2001. The neutral preference was 0.20
Favonigobius
gymnauchen Repomucenus spp. Tarphops oligolepis
Contents Mean (range) Mean (range) Mean (range)
Mysidae
Iiella ohshimai 0.303 (0.004–0.899) 0.242 (0–0.84) 0.156 (0.006–0.558)
Nipponomysis ornata 0.576 (0.083–1) 0.524 (0.247–0.732) 0.747 (0.241–1)
Anisomysis ijimai 0.402 (0.153–0.847) 0.460 (0.180–0.740) 0.203 (0–0.541)
Gammaridea 0.093 (0–0.217) 0.203 (0–1) 0.037 (0–0.178)
Crangon uritai. 0.221 (0–1) 0.150 (0–0.637) 0.015 (0–0.089)
Prey Predator values were very low. Newly settled Japanese floun-
Mysid
Goby der, however, was not found in the guts of the fish
Favonigobius gymnauchen
Iiella ohshimai examined and predation pressure on larval and
Dragnet
juvenile Japanese flounder by the three fish might
Repomucenus spp. not be high in the present study field.
Mysid
Nipponomysis ornata
Flounder
Tarphops oligolepis ACKNOWLEDGMENTS
Crangonid shrimp
Crangon uritai Japanese flounder We are grateful to Mr Y. Kameno, Ohama Fisheries
Paralichthys olivaceus
Cooperative Association, Mr Y. Kimura, Fukui
%IRI 5–10 : 10–30: 30–60: 60–: Prefectual University and the staff of the Kagawa
Overlap index 0.4–0.5: 0.5–0.6: 0.6–0.7: Prefectural Fisheries Experimental Station for
Fig. 4 Schematic diagram of food habits of Favonigo-
assistance in sampling. We thank Dr J. Shoji, Kyoto
bius gymnauchen, Repomucenus spp., Tarphops oligole- University and Dr Y. Natsukari, Nagasaki Univer-
pis and Paralichthys olivaceus. Diet overlaps and sity, for their useful information on feeding ecology
predator-prey relationship are shown. Data on the of fish. We are also grateful to Dr Y. Hanamura,
Japanese flounder after Yamamoto et al.23 National Research Institute of Fisheries and Envi-
ronment of Inland Sea, for useful advice on
crangonind shrimp. This work was partly sup-
ported by the Fisheries Agency of Japan. Thanks
crangonid shrimp) (Fig. 4). The mean body length are also due to the referees who made constructive
of mysids, crangonid shrimp and gammarids in the comments on the manuscript.
stomach of Japanese flounder collected in this
study area was 5.0 mm (range: 1.5–10.5 mm),
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