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164 ERIK BONSDORFF AND EA MARIA BLOMQVIST<br />

TABLE I11<br />

Fish species utilising <strong>the</strong> <strong>shallow</strong> <strong>water</strong> (0-5 m) archipelago areas of <strong>the</strong> nor<strong>the</strong>rn<br />

Baltic Sea, <strong>the</strong>ir ecological origin (fresh<strong>water</strong> = F, and marine = M), mode of<br />

feeding (predatory = Pr, planktivorous = P1, omnivorous = Om), and regi<strong>on</strong> of<br />

main occurrence (inner archipelago = I, middle archipelago = M, and outer<br />

archipelago = 0). Note that most fish species utilising <strong>the</strong> <strong>shallow</strong> <strong>water</strong>s also<br />

c<strong>on</strong>sume benthos, and thus participate in <strong>the</strong> benthic interacti<strong>on</strong>s of <strong>the</strong>se habitats<br />

(Blomqvist, 1982, 1984, 1986; Mattila & B<strong>on</strong>sdorff, 1988; B<strong>on</strong>sdorff & Blomqvist,<br />

1989; Rask, 1989; B<strong>on</strong>sdorff et al., 1990; Koli, 1990; Wistbacka, 1992).<br />

Species Origin Feeding type Occurrence<br />

-<br />

Pike (Esox lucius) F<br />

Whitefish (Coreg<strong>on</strong>us lavaretus) S. L. F<br />

Perch (Perca juviatilis) F<br />

Pikeperch (Stizostedi<strong>on</strong> lucioperca) F<br />

Ruffe (Gymnocephalus cernua) F<br />

Ide (Leuciscus idus) F<br />

Roach (Rurilus rurilus) F<br />

Rudd (Scardinius erythrophrhalmus) F<br />

White bream (Blicca bjoerkna) F<br />

Bream (Abramis brama) F<br />

Crucian carp (Carassius carassius) F<br />

Bleak (Albumus alburnus) F<br />

Cod (Gadus morhua) M<br />

Burbot (Lota lora) F<br />

Minnow (Phoxinus phoxinus) F<br />

Baltic herring (Clupea harengus membras) M<br />

Sprat (Sprartus spratrus) M<br />

Three-spined stickleback (Gasterosreus aculeatus) FIM<br />

Nine-spined stickleback (Pungitius pungitius) F<br />

Fifteen-spined stickleback (Spinachia spinachia) M<br />

Comm<strong>on</strong> goby (Pomatoschistus microps) M<br />

Sand goby (P. minutus) M<br />

Two-spotted goby (Gobiusculus flavescens) M<br />

Black goby (Gobius niger) M<br />

Eelpout (Zoarces viviparus) M<br />

Bullhead (C<strong>on</strong>us gobio) F<br />

Four-horned sculpin (Myoxocephalus quadricornis) M<br />

Sea scorpi<strong>on</strong> (Ceratocottus bubalis) F<br />

Flounder (Platicthys Jesus) M<br />

Turbot (Pserta maxima) M<br />

Deep-snouted pipefish (Syngnathus typhle) M<br />

Straight-nosed pipefish (Nerophis ophidi<strong>on</strong>) M<br />

Pr (fish)<br />

Pr (plankt<strong>on</strong>, benthos)<br />

Pr (benthos, fish)<br />

Pr (fish)<br />

Pr (benthos)<br />

Pr (benthos)<br />

Om<br />

Om<br />

Om<br />

Om<br />

Om<br />

PI<br />

Pr (benthos, fish)<br />

Pr (fish, benthos)<br />

Pr (benthos)<br />

PI<br />

PI<br />

PR (benthos, plankt<strong>on</strong>)<br />

Pr (benthos, plankt<strong>on</strong>)<br />

Pr (benthos, plankt<strong>on</strong>)<br />

Pr (benthos)<br />

Pr (benthos)<br />

Pr (benthos, plankt<strong>on</strong>)<br />

Pr (benthos)<br />

Pr (benthos)<br />

Pr (benthos, roe)<br />

Pr (benthos)<br />

Pr (benthos)<br />

Pr (benthos)<br />

Pr (benthos)<br />

PI<br />

PI<br />

1-0<br />

M-0<br />

1-0<br />

I+M<br />

I-tM<br />

I+M<br />

1-0<br />

I+M<br />

I-M<br />

I -- M<br />

I<br />

1-0<br />

0<br />

I (-' 0)<br />

M-0<br />

M+O<br />

0<br />

M--0<br />

1-0<br />

M--0<br />

I--M(0)<br />

(I) M - 0<br />

0<br />

1+0<br />

0<br />

M+O<br />

0<br />

0<br />

0<br />

0<br />

M+O<br />

M+O<br />

Mattila & B<strong>on</strong>sdorff, 1988; Nellbring, 1988; Aarnio & B<strong>on</strong>sdorff, 1992), and<br />

with changed size of <strong>the</strong> fish (Aarnio & B<strong>on</strong>sdorff, 1992; Mattila, 1992). Size<br />

selecti<strong>on</strong> within given prey populati<strong>on</strong>s (Le<strong>on</strong>ardss<strong>on</strong> et al., 1988) may ulti-<br />

mately participate in <strong>the</strong> structuring of <strong>the</strong> benthic community (B<strong>on</strong>sdorff et al.,<br />

1986b; Nellbring, 1988; Mattila, 1992). Many fish species are fur<strong>the</strong>rmore able<br />

to alter <strong>the</strong>ir feeding niche with changes in <strong>the</strong> envir<strong>on</strong>ment (for example vari-<br />

ati<strong>on</strong>s in <strong>water</strong> temperature, spatial heterogeneity etc.), or with altered com-<br />

petiti<strong>on</strong> for food (Magnhagen & Wiederholm, 1982; Bergman, 1987, 1988). If,<br />

<strong>on</strong> <strong>the</strong> o<strong>the</strong>r hand, <strong>the</strong> availability of food (i.e. <strong>the</strong> presence and abundance of<br />

zoobenthos) changes dramatically al<strong>on</strong>g a geographic gradient, this may regulate

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