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Ontogenetic Diet Shifts and Resource Partitioning Among Piscivorous Fishes in the Venezuelanllanos

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Ontogenetic Diet Shifts and Resource Partitioning Among Piscivorous Fishes in the Venezuelanllanos Environmental Biology of Fishes 26: 177-199. 1989. @ 1989 Klu~'er Academic Publishers. Printl'd in thl' Netherlands Ontogenetic diet shifts and resource partitioning among piscivorous fishes in the Venezuelanllanos Kirk O. Winemiller Departmentof Zoology, The Universityof Texas,Austin, TX 78712,U.S.A Received18.8.1988 Accepted12.12.1988 Key words: Caquetia, Charax. Diet breadth, Diffuse competition, Gymnotus, Hoplias. Resource overlap, Piranha,Pygocentrus, Rhamdia. Seasonality, Se"asalmus Synopsis Resource utilization by nine abundant piscivores from a diverse tropical fish assemblagewas examined over the course of a year. All nine speciesexhibited peak reproduction during the early wet seasonand a similar sequenceof size-dependent shifts from a diet composed primarily of microcrustacea, to aquatic insects, and finally fishes. Three piranha species specialized on fish firis, particularly at subadult size classes (SL 30-80mm). Gradual dessication of the floodplain during the transition season was associated with fish growth, increased fish density, and decreasedaquatic primary productivity and availability of invertebrate prey. Based on 118resource categories, averagepairwise diet overlap was low during all three seasons:wet, transition, and dry. Of 72 species pairings, only one pair of fin-nipping piranhas exhibited high overlap simultaneously on three niche dimensions: food type, food size, and habitat. Adults of two species, a gymnotid knifefish and pimelodid catfish, were largely nocturnal. Patterns of habitat utilization indicate that piranhas may restrict diurnal use of the open-water region by other piscivores. Collective diet overlap of individual piscivore specieswith the other eight feeding guild members and collective overlaps with the entire fish community each revealed two basic seasonal trends. Four species that showed an early switch to piscivory also showed a high degree of diet separation with both the guild and community at large on a year-round basis. The five remaining speciesshowed lowest collective diet overlaps during the transition seasonwhen availability of invertebrates was reduced and fish densities were maximal. Whereas predation may playa role in habitat separation, diffuse competition for food resources during the approximately four-month transition seasonprobably is the principal factor yielding patterns of diet specialization. Introduction bers provides the most straightforward means of attaining a preliminary assessmentof the relative Most diverse taxonomic groups of organisms ex- importance of interactive processeswithin natural hibit divergence in trophic biology (e .g., cichlid communities. Whereas descriptive studies that re- fishes, Greenwood 1981; ostariophysan fishes, veal patterns of resource subdivision are limited in Fink & Fink 1981).Whether or not ecomorpholog- their abilities to demonstrate the mechanismscre- ical divergence is random or adaptive in the context ating observed relationships (Toft 1985), thought- of community interactions can be debated (Strong ful comparisons and the weight of evidence derived et al. 1979, Grant 1981). The study of resource from numerous studies already have yielded much exploitation among syntopic feeding guild mem- insight (Schoener 1974, Helfman 1978, Sale 1979, 178 Werner 1979.Toft 1985).In a recentreview of the focuseson the piscivore guild as a potential unit for literature on fishes.Ross (1986) observed greatest competitive biological interactions. Given the cli- partitioning on food dimensions.followed by hab- mate and natural history of the Cano Maraca eco- itat and time. Many studiestreating two or more system, several basic questions can be asked: (1) nichedimensions have shown complementarity. in- Do piscivorous fishes partition food and habitat, dicatinga potential effect of interspecificcompeti- and if so, what is the influence of seasonalchanges tion. An interactive effect betweencompetition. in their environment? (2) What are the relation- predation, and temporal shifts in abiotic environ- ships between ontogeny, resource partitioning, mental parametershas been suggestedfor some and seasonal changes in resource availability? (3) systems(Werner & Hall 1977,Werner et al. 1983, Given seasonal patterns of resource availability Mittlebach 1984. and see Schoener1974. Toft and exploitation by consumers,is there evidence of 1985).Ross (1986)observed a generalpattern of interactive effects on piscivore feeding behavior? studiesrevealing less evidence of interactiveeffects (4) Do seasonal trends of pairwise species diet on communitystructure in harsherenvironments. overlaps follow patterns from collective diet over- Seasonalchanges in resourceavailability and on- laps within the piscivore guild or total fish commu- togeneticdiet shifts can affect both predator-prey nity, and if not, which index offers a better assess- and competitive interactionsamong size-structur- ment of the likelihood of interactive effects? ed fish populations. The present study explores seasonaland size-relatedpatterns of resourceex- ploitation within the piscivore guild of a diverse Methods neotropical fish community. Although varying greatlyin both duration and breadthof taxonomic Studysite comparison, several recent studies have shown markedresource partitioning amongtropical fishes During every month of 1984,fishes were collected (Angermeier& Karr 1983,Power 1983,Moyle & from Calio Maraca, a swamp-creek of the Rio Senanayake1984, Schut et al. 1984,Watson & Apure-Orinoco drainage in the western llanos of Balon 1984,Araujo-Lima et al. 1986,Hyslop 1986, Venezuela (8° 52' 30" Lat. N; 69°27' 40" Long. W). Prejs & Prejs 1987,Nico & Taphorn 1988).Fish The region studied (termed an 'estero') lies within populationsfrequently are size structureddue to low, flat terrain that experiences extensive sheet small hatching or neonate size and continuous flooding during the wettest months (late May-Au- growth. Body size significantly affects the size gust). During this time, a large region bordering rangeof food particlespotentially available to con- the creek channel is converted from exposed, sun- sumers(Brooks & Dodson 1965. Werner 1974, baked soil and thorn-scrub habitat into a produc- Magnan& Fitzgerald1984). Furthermore, size dis- tive marsh with diverse aquatic vegetation dom- tributions of fish populations directly influence inated by Cyperus sp., Eichhornia diversifolia, and predation rates on different ageclasses and often Heteranthera reniformis. During the driest months the spatial distribution of fishes (Fraser& Cern (Dec.-May), the aquatic habitat is reduced to a 1982,Mittlebach 1984,Power 1984,Werner & Gil- network of mud-bottom pools blanketed by Pistia liam 1984). stratiotes, Salvinia spp., and Lemna spp.. Dis- The studysite, Calio Maraca,experiences highly solved oxygen concentrations are reduced at this seasonalrainfall that is characteristicof manyparts time and many fishes rely on special respiratory of central and southern Venezuela.Most fishes adaptations for survival. Annual rainfall in the re- reproduceduring the early and peakwet season,a gion was 1300mm during 1984,with 860mm falling situationthat leadsto temporalchanges in average between June and September (Fig. 1). Based on body sizefor mostspecies (Winemiller, unpublish- changes exhibited by stream physico-chemical ed). Although similar data were collectedfor all parameters associated with rainfall and their ef- fishesthat occurred at the site, this investigation fects on aquatic organisms (Winemiller 1987a, 179 three wet months, collecting effort expended was approximately equal during each sampling period. More total time was spent collecting during the first ! three months of the wet seasondue to the compara- ~ tive inefficiency of adult fish captures in the newly- i.. 0 expanded aquatic environment. All aquatic hab- itats in the estero region were sampled for a mini- . .. ... .10. ,.. -0' A~, _0' .lu. .lul ... 00' Mo. 0.. mum of six hours (habitats combined), and a col- lecting outing was terminated when two hours of sampling yielded no additional species.Collections Fig. 1. Average monthly rainfall (closed symbols) and maximum used for comparisons of relative densities were midpool depth measured each month (open symbols) during the made during either one or two days between the study period at Cano Maraca. 11th and 28th of each month. Additional collec- tions were made on other dates to supplement stan- 1987b), the annual cycle was divided into three dard samples of uncommon species. A complete terms of equal duration: dry season (Jan.-Apr.), listing of collecting dates and sample sizes for pre- wet season (May-Aug.), and transition season served specimens appears in Winemiller (1987a). (Sept.-Dec.). Dry season conditions gave way to Collecting activities should not have affected nat- the wet season very abruptly with the commence- ural fish densities between sampling periods, be- ment of heavy rains, whereas the wet seasonslowly cause(1) only a fraction of the total aquatic habitat graded into the next dry season over a prolonged was sampled on a given date and (2) fishes could period (i.e., transition season). During the transi- move freely between the estero region and the tion season, fish densities increased and biotic in- pennanent channel downstream (except during teractions intensified within gradually diminishing March and April). All collected individuals were aquatic habitats in the floodplain. Estimates of fish preserved in 15% formalin rather than the usual biomassat Cano Maraca
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