A comparison of univariate and multivariate numerical and graphical techniques for determining inter- and intraspecific feeding relationships in estuarine fish

The advantages and disadvantages of various numerical and graphical techniques for the analysis of inter- and intraspecific feeding relationships of fishes were examined. All methods have been cited in the literature since 1988. The index of preponderance, the resultant index and graphical methods proposed by Costello and Tokeshi were used to illustrate the relative importance of prey species to an individual fish species with no differentiation between size classes. Inter- and intraspecific competition and niche overlap were determined from multivariate analysis [the ordination technique, detrended correspondence analysis, cluster analysis by the Bray-Curtis equation, per cent overlap and two-way indicator species analysis (TWINSPAN)]. The identity of the prey organisms are not lost in the comparisons, and the value of this is determined through comparison with techniques such as Shannon-Wiener which obscure these data. The Shannon-Wiener diversity index was combined with an analysis of 'evenness' to refine the technique further to assess niche breadth, as was the Levins index. The study shows that to give an estimate of competition within the community, it is important to assess the data with respect to seasonal and temporal patterns using multivariate analysis.     

Dietary niche breadth in a local community of passerine birds: an analysis using phylogenetic contrasts

The analysis of a local community of forest passerines (13 species) using phylogenetic contrasts shows a correlation between body size of bird species and mean prey size, minimum prey size, maximum prey size and the size range of dietary items. This suggests that larger birds drop small prey taxa from their prey list, because of the difficulty of capturing very small prey, for energetic reasons or because of microhabitat usage. We find some support for the third hypothesis. Dietary niche breadth calculated across prey taxa is not related to body size. Dietary niche breadth, however, is correlated with size-corrected measurements of the bill and locomotor apparatus. Long and slender bills increase the dietary niche breadth. Thus subtle differences constrain foraging and the techniques of extracting certain prey taxa form crevices. Dietary niche breadth and foraging diversity are positively correlated with population density: at least locally dietary generalists occur at higher breeding densities than specialists.     

Competition versus opportunism: Diet and trophic niche relationship of two sympatric apex carnivores in a tropical forest

There is a little understanding of how apex carnivores partition their diet to coexist. We studied food habit and trophic niche overlap of two apex carnivores, tiger and leopard in the tropical forest of Similipal tiger reserve, eastern India. We used line transect and scat analysis method, to estimate the prey availability and determine the diet and prey selection of two apex carnivores. Tigers consumed mostly large and medium-sized prey, whereas leopards mostly consumed medium and small-sized prey. Both carnivores were not random in their consumption of prey, sambar and wild pigs were selectively consumed by tigers, whereas leopards selectively consumed wild pigs, barking deer and mouse deer. Dietary overlap between two carnivore species was moderate (Pianka's niche overlap index: 0.55), and trophic niche breadth (Levin's standardized niche breadth) of the leopard (0.52) was greater than that of the tiger (0.37). Overall, tigers exhibited specialized feeding habits, whereas leopards showed generalist feeding habits. Our study highlights the opportunistic nature of leopards and probably a reason for the species successfully coexist with tigers.     

Trends in the prey size-based trophic niches of feral and House Cats Felis catus L.

House Cats Felis catus L., whether attached to human households or not, appear to be versatile opportunistic predators. Their principal prey in most areas are mammals (rodents and rabbits), with bird prey secondary. Trophic niche breadth, as measured by the standard deviation of the spectrum of logarithmically transformed prey sizes ('SLH'), shows a latitudinal trend, being greater in low latitudes: it is also greater in periods of high prey availability. This appears to be influenced by inclusion of very small prey, especially insects, in areas and seasons when they are available. Both the niche breadth and the mean prey size (niche position) appear to be constant as population mean cat size increases. The most common prey size for cats is about 1% of their own body weights, which is much less than most previously reported values for carnivores.     

Do lizard communities partition the trophic niche? A worldwide meta‐analysis using null models

Lizards have been widely used as model organisms in community ecology over the past 30 years. I have reviewed more than 50 studies from the literature on the community ecology of lizards worldwide, with a focus on studies from 1990–2007. I determined if there is support for the hypotheses that many lizard communities 1) are non‐randomly organized along the trophic niche dimension, and 2) partition the available food resource to minimize interspecific competition. I used data on number of prey items and percentage of prey volume, to calculate dietary overlap among species. I compared true datasets to randomly generate new datasets produced by 3×10⁴ Monte Carlo permutations using two algorithms (RA2 and RA3). The great majority (more than 80%) of the communities were randomly organized along the trophic niche dimension using RA2 or RA3; the frequency of occurrence of detecting a non‐random structure in lizard community dietary studies did not differ significantly between datasets based on either number of prey items or prey volume. Thus, lizard communities usually do not partition the trophic niche axis. Concerning the number of prey items, logistic regression models showed that the presence of a structure in the dataset did not depend on number of species, method employed to obtain dietary data, matrix size, or location (continent), but instead significantly depended on whether a community was tropical. Concerning prey volume, presence of a structure in the dataset also did not depend on number of species, method employed to obtain dietary data, and tropical vs non‐tropical origin, but was marginally dependent on continent (South America was favoured for identifying a structure in the dataset) and matrix size. In general, a structure in the dataset was more often uncovered by using RA2 than RA3. Overall, I conclude that 1) lizard communities are unlikely to partition available food resources, and 2) as the method for identifying dietary items does not allow for accurate prey identification (at least at the genus level), there is a risk of obtaining false null results about community structure. This is because structure along the trophic niche axis is genuinely rare in lizards and also because inappropriate methods may erroneously lead to this conclusion. Therefore, conclusions from studies utilizing stomach dissection as a source of diet data (still common in the literature) should be cautiously considered for ethical reasons and because it is difficult to find any difference between stomach‐dissection and fecal‐pellet data when assessing the presence of a structure in datasets. My data also showed there are minor differences in the probability of detecting a structure in datasets using prey item number or prey volume data for lizards. The biases, both statistical and biological, associated with this meta‐analysis are discussed.     

Quantifying Inter- and Intra-Population Niche Variability Using Hierarchical Bayesian Stable Isotope Mixing Models

Variability in resource use defines the width of a trophic niche occupied by a population. Intra-population variability in resource use may occur across hierarchical levels of population structure from individuals to subpopulations. Understanding how levels of population organization contribute to population niche width is critical to ecology and evolution. Here we describe a hierarchical stable isotope mixing model that can simultaneously estimate both the prey composition of a consumer diet and the diet variability among individuals and across levels of population organization. By explicitly estimating variance components for multiple scales, the model can deconstruct the niche width of a consumer population into relevant levels of population structure. We apply this new approach to stable isotope data from a population of gray wolves from coastal British Columbia, and show support for extensive intra-population niche variability among individuals, social groups, and geographically isolated subpopulations. The analytic method we describe improves mixing models by accounting for diet variability, and improves isotope niche width analysis by quantitatively assessing the contribution of levels of organization to the niche width of a population.     

Sexual Dimorphism of Head Size in Phrynocephalus przewalskii: Testing the Food Niche Divergence Hypothesis

Sexual size dimorphism （SSD） is a general phenomenon in lizards, and can evolve through sexual selection or natural selection. But natural selection, which was thought to operate mainly through reducing the competition be- tween the two sexes （niche divergence hypothesis）, gave rise to a lot of controversy. We tested the niche divergence hypothesis in the toad-headed lizard Phrynocephalus przewalskii by comparing diet composition and prey sizes between males and females. The species was found to be sexual dimorphic, with males having relatively larger snout-vent length, head width, head length, and tail length, while females have relatively larger abdomen length. Based on analysis of 93 studied stomachs, a total of 1359 prey items were identified. The most common prey items were formicid, lygaeid and tenebrionid. The two sexes did not differ in the relative proportions of prey size categories they consumed and the dietary overlap based on prey species was high （O = 0.989）. In addition, the meal size, the volume or any maximal dimension of the largest prey item in the stomach was not explained by the sexes. According to our results, food niche divergence might not play an important role in the SSD evolution ofP. przewalskii.     

Generalist niche, specialist strategy: the diet of an Australian percichthyid

Dietary analysis revealed that an impoundment population of Australian bass Macquaria novemaculeata holds a generalist niche, but one arising from persistent individual specialization and interindividual variation. This 'individual specialist' strategy appeared adaptive, but the strength of individual specialization was largely independent of variation in diet composition, except during blooms of Daphnia sp. Diet composition and dietary overlap showed only moderate ontogenetic variation, and niche breadth showed no relationship with ontogeny. Macquaria novemaculeata showed an asymmetric predator and prey size distribution, consistent with many aquatic predators, with positive relationships between fish size and average, maximum and minimum prey size. There was no asymmetry in the relative size-based niche breadths of individuals, however, which indicates that the niche is a fixed 'window' of relative prey sizes. The difference in the dietary niche and prey-size relationships of M. novemaculeata at the population and individual levels highlights the necessity of assessing the niche at both these levels.     

Complex interactions between components of individual prey specialization affect mechanisms of niche variation in a grasshopper-hunting wasp

1.?Individual foraging behaviour defines the use of resources by a given population and its variation in different ways such as, for example, unpredictable interactions between taxon-biased and size-biased selection. Here we investigated how the environmental availability of prey and individual specialization, for both prey taxa and prey size, shape niche variation across generations in the grasshopper-hunting digger wasp Stizus continuus. 2.?The population of S.?continuus expressed selective predation, females mainly hunting species encountered on large bushes; diet changed across generations, due more to size increase in potential prey than to changes in the orthopteran community. 3.?Individual females of both generations weakly overlapped the size and taxa of prey, and the niche width of the second generation increased for both prey size and taxa. 4.?The greater variance in prey size in the environment accounted for the enlarged prey size niche of the second generation, but the load-lifting constraints of the wasps maintained individual prey size specialization constant. In contrast, the enlarged prey taxon niche paralleled a smaller overlap of diets between wasps in the second generation. 5.?Increased niche width in the S.?continuus population was thus achieved in two ways. Regarding prey size, all individuals shifted towards the use of the full set of available resources (parallel release). For prey taxa, according to the classical niche variation hypothesis, individuals diverged to minimize resource use overlap and perhaps intraspecific competition. These two mechanisms were observed for the first time simultaneously in a single predator population.     

Trophic niche partitioning and diet composition of sympatric fin (Balaenoptera physalus) and humpback whales (Megaptera novaeangliae) in the Gulf of Alaska revealed through stable isotope analysis

Fin and humpback whales are large consumers that are often sympatric, effectively sharing or partitioning their use of habitat and prey resources. Stable carbon and nitrogen isotopes in the skin of fin and humpback whales from two regions in the western Gulf of Alaska, Kodiak, and Shumagin Islands, were analyzed to test the hypothesis that these sympatric baleen whales exhibit trophic niche partitioning within these regions. Standard ellipse areas, estimated using Bayesian inference, suggested that niche partitioning between species is occurring in the Kodiak region but not in the Shumagin Islands. Isotopic mixing models based on stable isotopes from whales and local prey samples, were used to estimate possible diet solutions for whales in the Kodiak region. Comparison of isotopic niches and diet models support niche partitioning, with fin whales foraging primarily on zooplankton and humpback whales foraging on zooplankton and small forage fish. The results of this study show that niche partitioning between sympatric species can vary by region and may be the result of prey availability, prey preferences, or both.     

The effects of morphology,phylogeny and prey availability on trophic resource partitioning in an anuran community

Several factors influence the partitioning of trophic resources in ecological communities, such as morphology, evolutionary history, and resource availability. Although the effects of morphology, phylogeny, and resource availability on trophic ecology have long been explored by theoretical studies, little has been done to empirically test these relationships. Here, we tested whether phylogenetic and morphological distances correlate with trophic niche overlap using a path analysis of multiple partial regression of distance matrices. Also, we tested whether niche breadth is influenced by body size using Phylogenetic Generalized Least Squares analysis. Trophic niche overlap was better explained by morphology per se than by the phylogenetic distance. We also found that predator's body size influences niche breadth calculated considering prey traits and availability, but not when we do not include these availability data. Additionally, trophic niche breadth was usually smaller when we considered prey traits and availability, differently from niche overlap, whose values increased when we did not consider these data. Our findings show that the interpretation of trophic niche in communities changes if we consider availability data, affecting inferences about coexistence and trophic specialization. Our study contributes to understanding trophic specialization and emphasizes the importance of incorporating prey availability and their traits into diet analysis.     

Assessing niche partitioning of co‐occurring sibling bat species by DNA metabarcoding

Niche partitioning through foraging is a mechanism likely involved in facilitating the coexistence of ecologically similar and co‐occurring animal species by separating their use of resources. Yet, this mechanism is not well understood in flying insectivorous animals. This is particularly true of bats, where many ecologically similar or cryptic species coexist. The detailed analysis of the foraging niche in sympatric, cryptic sibling species provides an excellent framework to disentangle the role of specific niche factors likely involved in facilitating coexistence. We used DNA metabarcoding to determine the prey species consumed by a population of sympatric sibling Rhinolophus euryale and Rhinolophus mehelyi whose use of habitat in both sympatric and allopatric ranges has been well established through radio tracking. Although some subtle dietary differences exist in prey species composition, the diet of both bats greatly overlapped (O_jk = 0.83) due to the consumption of the same common and widespread moths. Those dietary differences we did detect might be related to divergences in prey availabilities among foraging habitats, which prior radio tracking on the same population showed are differentially used and selected when both species co‐occur. This minor dietary segregation in sympatry may be the result of foraging on the same prey‐types and could contribute to reduce potential competitive interactions (e.g., for prey, acoustic space). Our results highlight the need to evaluate the spatial niche dimension in mediating the co‐occurrence of similar insectivorous bat species, a niche factor likely involved in processes of bat species coexistence.     

Is dietary niche breadth linked to morphology and performance in Sandveld lizards Nucras (Sauria: Lacertidae)?

The functional characteristics of prey items (such as hardness and evasiveness) have been linked with cranial morphology and performance in vertebrates. In lizards particularly, species with more robust crania generally feed on harder prey items and possess a greater bite force, whereas those that prey on evasive prey typically have longer snouts. However, the link between dietary niche breadth, morphology, and performance has not been explicitly investigated in lizards. The southern African genus Nucras was used to investigate this link because the species exhibit differing niche breadth values and dietary compositions. A phylogeny for the genus was established using mitochondrial and nuclear markers, and morphological clusters were identified. Dietary data of five Nucras species, as reported previously, were used in correlation analyses between cranial shape (quantified using geometric morphometrics) and dietary niche breadth, and the proportion of hard prey taken and bite force capacity. Dietary niche breadth and the proportion of hard prey eaten were significantly related to cranial shape, although not once phylogeny was accounted for using a phylogenetic generalized least squares regression. The proportion of evasive prey eaten was a significant predictor of forelimb length when phylogeny was taken into account. We conclude that, in Nucras, the percentage of evasive prey taken co‐evolves with forelimb morphology, and dietary niche breadth co‐evolves with cranial shape. However, although head width is correlated with the proportion of hard prey eaten, this appears to be the result of shared ancestry rather than adaptive evolution. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 674–688.     

Importance of niche quality for Yellowhammer Emberiza citrinella nestling survival, development and body condition in its native and exotic ranges: the role of diet

The niche hypothesis predicts that some introduced species establish and spread successfully because their new environment provides expanded niche opportunities compared with their native environments. By investigating nestling survival, growth and body condition in relation to diet composition and prey abundance, we tested the prediction that the success of Yellowhammers Emberiza citrinella in New Zealand could be explained by the availability of better quality food resources in its introduced range compared with its native range. We found that Yellowhammer nestlings in New Zealand were larger and heavier than those in Britain, but that nestling survival and body condition did not differ between countries. The preferred prey items, which had some influence on nestling development and survival, were more prevalent in the diet of nestlings in New Zealand, suggesting that nestlings in the introduced range had a superior quality diet. However, there was little evidence to support the prediction that the preferred prey were more abundant in the introduced range, implying that some other factor such as prey accessibility or adult fitness may account for the differences in diet between countries.     

Seasonal variation in terrestrial resource subsidies influences trophic niche width and overlap in two aquatic snake species: a stable isotope approach

Quantifying diet is essential for understanding the functional role of species with regard to energy processing, transfer, and storage within ecosystems. Recently, variance structure in the stable isotope composition of consumer tissues has been touted as a robust tool for quantifying trophic niche width, a task that has previously proven difficult due to bias in direct dietary analyses and difficulties in integrating diet composition over time. We used carbon and nitrogen stable isotope analyses to examine trophic niche width of two sympatric aquatic snakes, banded watersnakes Nerodia fasciata and black swamp snakes Seminatrix pygaea inhabiting an isolated wetland where seasonal migrations of amphibian prey cause dramatic shifts in resource availability. Specifically, we characterized snake and prey isotope compositions through time, space, and ontogeny and examined isotope values in relation to prey availability and snake diets assessed by gut content analysis. We determined that prey cluster into functional groups based on similarity of isotopic composition and seasonal availability. Isotope variance structure indicated that the trophic niche width of the banded watersnake was broader (more generalist) than that of the black swamp snake. Banded watersnakes also exhibited seasonal variation in isotope composition, suggesting seasonal diet shifts that track amphibian prey availability. Conversely, black swamp snakes exhibited little seasonal variation but displayed strong ontogenetic shifts in carbon and nitrogen isotope composition that closely paralleled ontogenetic shifts in their primary prey, paedomorphic mole salamanders Ambystoma talpoideum. Although niche dimensions are often treated as static, our results demonstrate that seasonal shifts in niche dimensions can lead to changes in niche overlap between sympatric species. Such short‐term fluctuations in niche overlap can influence competitive interactions and consequently the composition and dynamics of communities and ecosystems.     

Diet of coastal foraging Eurasian otters (<Emphasis Type="Italic">Lutra lutra</Emphasis> L.) in Pembrokeshire south-west Wales

The importance of the marine environment to Eurasian otters is currently poorly understood. Wales is one of the few countries where coastal activity has been recorded and an increase in marine otter sightings could indicate remarkable developments within Welsh populations. The trophic niche of coastal otter populations around Pembrokeshire was investigated over a 12-month period. Marine activity was more widespread than previously thought and marine prey formed the largest component of otter diet, although, otters also consumed freshwater and terrestrial prey throughout the year. Otter diet was very diverse compared to other European coastal populations and a spring contraction in trophic niche width coincided with the estimated timing of breeding activity. Seasonal variation in prey composition was predominantly due to differences in the consumption of alternate prey types. In areas where wetlands are fragmented and populations of freshwater fish are declining, the marine environment may become an increasingly important habitat for otters. It is necessary to define the historical importance of coastal populations to otter conservation. Coastal areas are often subject to pressure from human activities, so the impact of disturbance needs to be assessed. Importantly, there is no verified otter survey method for coastal areas, so the use of marine habitat is likely to be underestimated.     

Ecologically based definition of seasons clarifies predator–prey interactions

Species interactions within food webs are driven by multiple constraints, including those imposed by seasonal changes in the environment. Ecologically sound definitions of seasons may therefore be a prerequisite for clarifying predator prey interactions. Most studies define biological seasons based on fixed schedules or on temporal changes in a single movement measurement. We used a novel clustering approach based on homogeneous space‐use patterns of GPS‐collared animals to reveal 7 biological seasons for caribou Rangifer tarandus caribou, and 5 for both moose Alces alces and grey wolves Canis lupus interacting in a boreal ecosystem. Subsequent evaluation of niche overlap showed that, as predicted, wolves had a stronger spatio‐temporal connection with moose, its main prey, than with caribou. Movement constraints and limiting resource distributions similarly affected all species in some instances, but also caused temporal changes in the extent of niche overlap between wolves and its two prey. The risk that caribou faced was not only linked to the niche overlap with wolves, but also to the extent of wolf‐moose niche overlap during the same period. Food‐web properties emerged from the analysis, with temporal changes in relative niche overlap reflecting the strength of trophic interactions during the year. Our study demonstrates how the study of trophic interactions can benefit from comprehensive definitions of biological seasons.     

Feeding biology of the flatfish Citharichthys spilopterus (Bothidae) in a tropical estuary of Mexico

The flatfish Citharichthys spilopterus Günther 1862 is the most common bothid in the coastal lagoons of the Gulf of Mexico. The objective of the present study was to describe the trophic biology of this species in the tropical coastal lagoon of Tampamachoco, Mexico. For the diet analysis, we used multivariate discriminant analysis and trophic niche breadth. The morphological analysis showed that the features of the bucco-pharyngeal cavity, the large stomach and the short intestine of this flatfish relate to a carnivorous habit. The overall diet showed that this species is a third-order consumer, feeding mainly on fish (52%, mainly gobies) and crustaceans (36%, mostly decapods). Diet and trophic niche breadth showed no significant differences between sexes (P > 0.1). By contrast, there were significant ontogenetic differences in the diet and trophic niche breadth, where larger C. spilopterus were almost entirely piscivorous and showed the narrowest trophic niche. As the flatfish grew in size there was a trend toward the consumption of larger prey. The importance of copepods and peracarids correlated inversely with flatfish size (P < 0.001), but fish prey correlated directly to flatfish size (P < 0.001). Likewise, there was an inverse significant correlation between niche trophic breadth and flatfish size (P < 0.005). There were also significant seasonal differences in the diet (P < 0.02), related to the availability and vulnerability of prey in the lagoon during the dry and rainy seasons. However, the trophic niche showed no significant differences between seasons. Finally, we discuss the advantages of discriminant analysis applied to evaluate differences among diets of fish groups when compared with other bivariate and multivariate techniques.     

Sensory-based niche partitioning in a multiple predator–multiple prey community

Many predators and parasites eavesdrop on the communication signals of their prey. Eavesdropping is typically studied as dyadic predator–prey species interactions; yet in nature, most predators target multiple prey species and most prey must evade multiple predator species. The impact of predator communities on prey signal evolution is not well understood. Predators could converge in their preferences for conspicuous signal properties, generating competition among predators and natural selection on particular prey signal features. Alternatively, predator species could vary in their preferences for prey signal properties, resulting in sensory-based niche partitioning of prey resources. In the Neotropics, many substrate-gleaning bats use the mate-attraction songs of male katydids to locate them as prey. We studied mechanisms of niche partitioning in four substrate-gleaning bat species and found they are similar in morphology, echolocation signal design and prey-handling ability, but each species preferred different acoustic features of male song in 12 sympatric katydid species. This divergence in predator preference probably contributes to the coexistence of many substrate-gleaning bat species in the Neotropics, and the substantial diversity in the mate-attraction signals of katydids. Our results provide insight into how multiple eavesdropping predator species might influence prey signal evolution through sensory-based niche partitioning.     

Species‐level predation network uncovers high prey specificity in a Neotropical army ant community

Army ants are among the top arthropod predators and considered keystone species in tropical ecosystems. During daily mass raids with many thousand workers, army ants hunt live prey, likely exerting strong top‐down control on prey species. Many tropical sites exhibit a high army ant species diversity (>20 species), suggesting that sympatric species partition the available prey niches. However, whether and to what extent this is achieved has not been intensively studied yet. We therefore conducted a large‐scale diet survey of a community of surface‐raiding army ants at La Selva Biological Station in Costa Rica. We systematically collected 3,262 prey items from eleven army ant species (genera Eciton, Nomamyrmex and Neivamyrmex). Prey items were classified as ant prey or non‐ant prey. The prey nearly exclusively consisted of other ants (98%), and most booty was ant brood (87%). Using morphological characters and DNA barcoding, we identified a total of 1,103 ant prey specimens to the species level. One hundred twenty‐nine ant species were detected among the army ant prey, representing about 30% of the known local ant diversity. Using weighted bipartite network analyses, we show that prey specialization in army ants is unexpectedly high and prey niche overlap very small. Besides food niche differentiation, we uncovered a spatiotemporal niche differentiation in army ant raid activity. We discuss competition‐driven multidimensional niche differentiation and predator–prey arms races as possible mechanisms underlying prey specialization in army ants. By combining systematic prey sampling with species‐level prey identification and network analyses, our integrative approach can guide future research by portraying how predator–prey interactions in complex communities can be reliably studied, even in cases where morphological prey identification is infeasible.