Stable isotope ratios (typically of carbon and nitrogen) provide one representation of an organism's trophic niche and are widely used to examine aspects of food web structure. Yet stable isotopes have not been applied to quantitatively characterize community-wide aspects of trophic structure (i.e., at the level of an entire food web). We propose quantitative metrics that can be used to this end, drawing on similar approaches from ecomorphology research. For example, the convex hull area occupied by species in delta13C-delta15N niche space is a representation of the total extent of trophic diversity within a food web, whereas mean nearest neighbor distance among all species pairs is a measure of species packing within trophic niche space. To facilitate discussion of opportunities and limitations of the metrics, we provide empirical and conceptual examples drawn from Bahamian tidal creek food webs. These examples illustrate how this methodology can be used to quantify trophic diversity and trophic redundancy in food webs, as well as to link individual species to characteristics of the food web in which they are embedded. Building from extensive applications of stable isotope ratios by ecologists, the community-wide metrics may provide a new perspective on food web structure, function, and dynamics.
Genera within the South American cichlid tribe Geophagini display specialized feeding and reproductive strategies, with some taxa specialized for both substrate-sifting and mouth brooding. Several lineages within the clade also possess an epibranchial lobe (EBL), a unique pharyngeal structure that has been proposed to have a function in feeding and/or mouth brooding. A recently published genus-level phylogeny of Neotropical cichlids was used as the evolutionary framework for investigating the evolution of morphological features presumably correlated with diet and mouth brooding in the tribe Geophagini. We tested for possible associations between the geophagine epibranchial lobe and benthic feeding and mouth brooding. We also addressed whether the EBL may be associated with unique patterns of diversification in certain geophagine clades. Tests of binary character correlations revealed the EBL was significantly associated with mouth brooding. We also tested for a relationship between diet and morphology. We analyzed stomach contents and morphometric variation among 21 species, with data for two additional species obtained from the literature. Principal Components Analysis revealed axes of morphological variation significantly correlated with piscivory and benthivory, and both morphology and diet were significantly associated with phylogeny. These results suggest that the EBL could be an adaptation for either feeding or mouth brooding. The EBL, however, was not associated with species richness or accelerated rates of phyletic diversification.
Despite divergent evolutionary histories, Neotropical cichlids (Cichlidae) and Nearctic sunfishes (Centrarchidae) appear to have similar functional morphotypes and occupy similar ecological niches. We applied an integrative approach analyzing morphological traits, stomach contents, and stable isotope ratios (δ13C, δ15N) to investigate whether local assemblages of cichlids (Venezuela, Peru) and centrarchids (Texas) reveal one‐to‐one patterns of morphological and ecological convergence. Multivariate ordinations performed on diet and morphology datasets identified a broad overlap between cichlid and centrarchid assemblages. The functional morphology of the two groups has diversified in a convergent manner within the confines of ram‐suction modes of prey ingestion. Both groups had the same set of ecomorph types that corresponded to the same trophic niches, including substrate‐sifting invertivores, epibenthic invertebrate gleaners, and piscivores; the one exception was a molluscivorous sunfish, comprising a niche that was not represented in the two cichlid assemblages. Estimates of trophic positions based on stable isotope analysis revealed convergent vertical trophic structure; with few exceptions, fishes with similar morphologies had similar trophic positions. Large‐bodied piscivores had highest trophic positions, whereas small and medium‐bodied generalists and invertivores had low to intermediate trophic positions. Consistent patterns of ecomorphological convergence in these two perciform groups provide strong evidence for adaptation involving constrains in functional morphology associated with feeding. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109, 146–164.
The frogs of the family Ceratophryidae (comprised of three genera containing 12 species) are well known for their voracious feeding behavior and unique morphologies in both larval and post-metamorphic life stages. Nevertheless, relatively little is known about the diet of these species, especially in nature. We summarized the current knowledge of the diet of larval and post-metamorphic ceratophryids in both natural and experimental settings by conducting a literature review, specimen dissections, and field observations. Prior to this study, diet information did not exist for one fourth of the species in the family: Ceratophrys calcarata, Ceratophrys joazeirensis Ceratophrys stolzmanni, and Ceratophrys testudo. We add, for the first time, information on the diet of Ceratophrys calcarata. In addition, our survey revealed that relatively few detailed studies with large sample sizes have been conducted on the diet of these frogs. Frequently, knowledge of the diet of a species is derived from palatability experiments using captive individuals (often a single individual). From those diet studies that were conducted under natural conditions, ceratophryids can be classified as generalist, opportunistic predators, often preying on a wide variety of invertebrate and vertebrate prey. All species of ceratophryids for which we obtained information on diet were found to consume vertebrates, with anuran prey occurring in nearly all species. Even less is known about the diet of the tadpoles of these species; while cannibalism is assumed to be widespread, we found evidence for cannibalism at the larval stage in only three species. Future studies should attempt to understand the trophic ecology of these species under natural scenarios at both life stages, and an ontogenetic approach using individuals across multiple size classes could elucidate the niche shifts associated with these species from larva to adult.
Community assembly is affected by environmental filtering that restricts viable phenotypes and by species interactions that impose limits on interspecific trait similarity.Relative influences of these processes should vary according to habitat features and dispersal. Species dispersion within assemblage trait space also should vary in relation to species richness, strength of competition, and the spatiotemporal scale of analysis. We examined ecomorphological diversity of two freshwater fish families (Neotropical Cichlidae, Nearctic Centrarchidae) to test theories of local assembly from regional species pools and theories of species packing within mesohabitat patches. Cichlid and centrarchid assemblages were surveyed in four floodplain rivers (two in South America and two in North America) during low-water periods when fish densities are highest. Surveys were conducted in four mesohabitat types (submerged wood, leaf litter, rocks, sand bank) within river channels and floodplain lakes. We measured 23 morphological traits associated with locomotion and feeding. Principal components analysis was performed on the species 3 traits matrix, and species axis scores were used to calculate species pairwise Euclidean distances and indices of dispersion within assemblage morphospace: mean nearest-neighbor distance (indicating similarity), mean distance to centroid (assemblage morphospace size), and standard deviation of nearestneighbor distance (evenness of dispersion within assemblage morphospace). A null model was used to assess whether patterns were significantly nonrandom. When data for all mesohabitat types were combined for each river, species were significantly overdispersed and the assemblage morphospace was larger than predicted at random in every case. Analysis of assemblages within mesohabitat patches of different types revealed, in every case, significant overdispersion of species in morphospace indicative of limiting similarity. The total assemblage morphospace was greater than expected for tropical cichlids, but not for temperate centrarchids. Trends of species dispersion with assemblage morphospace in relation to species richness within mesohabitat patches were not consistent among or within river systems, possibly indicating that patches were already saturated with these perciform fishes. Interregional comparisons suggest an influence from both adaptive diversification and environmental filtering at broad spatial scales. At the scale of mesohabitat patches in lowland rivers, cichlid and centrarchid assemblages revealed patterns of trait complementarity that imply limiting similarity and strong influence of biotic interactions.
Montaña CG, Winemiller KO. Local‐scale habitat influences morphological diversity of species assemblages of cichlid fishes in a tropical floodplain river. Ecology of Freshwater Fish 2010: 19: 216–227. © 2010 John Wiley & Sons A/S Abstract – We examined the taxonomic and morphological diversity of cichlid fish assemblages in a floodplain river in Venezuela during the dry season at two spatial scales: macrohabitats (lagoons, main channels and creeks) and mesohabitats (leaf litter, sand banks, rocky shoals and woody debris). Nonmetric multidimensional scaling did not reveal differences for species assemblages among macro and mesohabitats. The first two axes from canonical correspondence analysis based on 19 species and six physical variables modelled >61% of the taxonomic variation in assemblages from rock shoals and woody debris, and 55% of variation in assemblages from sand banks and leaf litter. Principal components analysis based on 22 morphological variables yielded two dominant axes that explained >86% of variation in the cichlid assemblages. Morphological diversity was analysed to test the idea that assemblage structure is nonrandom, with structurally complex habitats supporting more species with more functional morphological diversity than simple habitats. Average and standard deviation (SD) of the morphological Euclidean distances of local assemblages among mesohabitats tended to decrease or be constant as the number of species increased. Regressions of the average nearest neighbour distance (NND) and SD of NND with species richness resulted in low and negative slopes of species assemblages among mesohabitats. These relationships suggest that when more species are added to a habitat patch, assemblage morphospace remains approximately constant, species average similarity increases and species dispersion in morphological space becomes more uniform. Results support that cichlids partition habitat at the local scale but not at the macrohabitat scale.
The Casiquiare River is a unique biogeographic corridor between the Orinoco and Amazonas basins. We investigated the importance of this connection for Neotropical fishes using peacock cichlids (Cichla spp.) as a model system. We tested whether the Casiquiare provides a conduit for gene flow between contemporary populations, and investigated the origin of biogeographic distributions that span the Casiquiare. Using sequences from the mitochondrial control region of three focal species (C. temensis, C. monoculus, and C. orinocensis) whose distributions include the Amazonas, Orinoco, and Casiquiare, we constructed maximum likelihood phylograms of haplotypes and analyzed the populations under an isolation-with-migration coalescent model. Our analyses suggest that populations of all three species have experienced some degree of gene flow via the Casiquiare. We also generated a mitochondrial genealogy of all Cichla species using >2000 bp and performed a dispersal-vicariance analysis (DIVA) to reconstruct the historical biogeography of the genus. This analysis, when combined with the intraspecific results, supports two instances of dispersal from the Amazonas to the Orinoco. Thus, our results support the idea that the Casiquiare connection is important across temporal scales, facilitating both gene flow and the dispersal and range expansion of species.
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