Optimal foraging constrains macroecological patterns: body size and dietary niche breadth in lizards

Aim  To explore and identify probable mechanisms contributing to the relationships among body size, dietary niche breadth and mean, minimum, maximum and range of prey size in predaceous lizards.
Location  Our data set includes species from tropical rainforests, semi-arid regions of Brazil, and from deserts of the south-western United States, Australia and the Kalahari of Africa.
Methods  We calculated phylogenetic and non-phylogenetic regressions among predator body size, dietary breath and various prey size measures.
Results  We found a negative association between body size and dietary niche breadth in 159 lizard species sampled across most evolutionary lineages of squamate reptiles and across major continents and habitats. We also show that mean, minimum, maximum and range of prey size were positively associated with body size.
Main conclusions  Our results suggest not only that larger lizards tend to eat larger prey, but in doing so offset their use of smaller prey. Reduction of dietary niche breadth with increased body size in these lizards suggests that large predators target large and more profitable prey. Consequently, the negative association between body size and niche breadth in predators most likely results from optimal foraging. Though this result may appear paradoxical and runs counter to previous studies, resources for predators may be predictably more limited than resources for herbivores, thus driving selection for more profitable prey.     

Patterns of prey size and taxonomic composition in larval fish: are there general size-dependent models?

The allometric relationship of maxilla length to larval fish weight approached a slope of 1/3 for 11 species from Conception Bay, Newfoundland. In 10 of these species, mean prey size increased with length of the maxilla but not constantly. In six species, niche breadth increased with length of maxilla. The rate of change in niche breadth was inversely related to the rate of change in mean prey size. Nauplii and copepodites of calanoids were selected positively, and cyclopoids negatively for almost all species of larval fish.     

Diet breadth variability in larval blue whiting as a response to plankton size structure

Diet breadth (measured as the S.D. of the log of prey size per larvae; SLH) of blue whiting micromesistius poutassou larvae followed a quadratic equation with larval size. In small larvae, diet breadth in terms of size (SLH), the mean and the maximum of the log of prey size per larvae (MLH and XLH, respectively) increased with larval size as prey size selection shifted to larger prey. In contrast, large larvae tended to reduce diet breadth of prey sizes ingested, focusing on the larger prey that were abundant, instead of raising the upper limit of prey sizes because of the low abundance of larger prey. Except for larvae at the onset of first feeding, number of prey stayed constant or decreased in relation to larval size. Both patterns (in small and large larvae) maintained a constant rate of increase of gut carbon content with increase in larval size. Large larvae appear to maintain the increase in gut carbon content during ontogenetic development by reducing diet breadth (SLH) and increasing selection towards the larger prey that are abundant.     

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.     

Niche dynamics of sympatric non-breeding shearwaters under varying prey availability

Variation in prey availability can cause changes in species interactions among marine predators. Foraging theory predicts that niche breadth will expand when resources become limited, possibly leading to higher niche overlap among sympatric species; however, a species’ niche can become constrained by interactions with other similar species, resulting in an inability to shift niche breadth or position. In coastal Newfoundland, Capelin Mallotus villosus is the main forage fish species and its availability (i.e. biomass) during the summer has varied considerably following a population collapse in the 1990s. During the summer, non-breeding Great and Sooty Shearwaters Ardenna gravis and A. grisea migrate and aggregate at annually persistent Capelin spawning sites. We used stable isotope ratios (δ¹³C, δ¹⁵N) of blood components (plasma, red blood cells) to investigate variation in isotopic niche breadth (95% prediction ellipse areas) and overlap of the two shearwater species during 2014, 2015 and 2016. Capelin availability varied among years, illustrated by lower peak biomass in 2015 (0.126 g/m²) and 2016 (0.027 g/m²) relative to 2014 (0.254 g/m²). The isotopic niche breadth (plasma) of both shearwater species expanded similarly from 2014 (0.65–0.66‰²) to 2015 (2.22–2.57‰²) and 2016 (1.15–1.42‰²), suggesting the incorporation of alternative prey types into their diet during years of lower prey availability. Isotopic niche overlap between Great and Sooty Shearwaters remained high across years (44–63%), however, providing little evidence for dietary niche partitioning during years of lower prey availability. Findings suggest that both shearwater species are flexible foragers and can modify their diet during the non-breeding season to accommodate fluctuations in prey availability.     

The influence of coastal access on isotope variation in Icelandic arctic foxes

To quantify the ecological effects of predator populations, it is important to evaluate how population-level specializations are dictated by intra- versus inter-individual dietary variation. Coastal habitats contain prey from the terrestrial biome, the marine biome and prey confined to the coastal region. Such habitats have therefore been suggested to better support predator populations compared to habitats without coastal access. We used stable isotope data on a small generalist predator, the arctic fox, to infer dietary strategies between adult and juvenile individuals with and without coastal access on Iceland. Our results suggest that foxes in coastal habitats exhibited a broader isotope niche breadth compared to foxes in inland habitats. This broader niche was related to a greater diversity of individual strategies rather than to a uniform increase in individual niche breadth or by individuals retaining their specialization but increasing their niche differentiation. Juveniles in coastal habitats exhibited a narrower isotope niche breadth compared to both adults and juveniles in inland habitats, and juveniles in inland habitats inhabited a lower proportion of their total isotope niche compared to adults and juveniles from coastal habitats. Juveniles in both habitats exhibited lower intra-individual variation compared to adults. Based on these results, we suggest that foxes in both habitats were highly selective with respect to the resources they used to feed offspring, but that foxes in coastal habitats preferentially utilized marine resources for this purpose. We stress that coastal habitats should be regarded as high priority areas for conservation of generalist predators as they appear to offer a wide variety of dietary options that allow for greater flexibility in dietary strategies.     

Only the largest terrestrial carnivores increase their dietary breadth with increasing prey richness

1. Animals should adapt their foraging habits, changing their dietary breadth in response to variation in the richness and availability of food resources. Understanding how species modify their dietary breadth according to variation in resource richness would support predictions of their responses to environmental changes that alter prey communities.
2. We evaluated relationships between the dietary breadth of large terrestrial carnivores and the local richness of large prey (defined as the number of species). We tested alternative predictions suggested by ecological and evolutionary theories: with increasing prey richness, species would (1) show a more diverse diet, thus broadening their dietary breadth, or (2) narrow their dietary breadth, indicating specialisation on a smaller number of prey.
3. We collated data from 505 studies of the diets of 12 species of large terrestrial mammalian carnivores to model relationships between two indices of dietary breadth and local prey richness.
4. For the majority of species, we found no evidence for narrowing dietary breadth (i.e. increased specialisation) with increasing prey richness. Although the snow leopard and the dhole appeared to use a lower number of large prey species with increasing prey richness, larger sample sizes are needed to support this result.
5. With increasing prey richness, the five largest carnivores (puma Puma concolor, spotted hyaena Crocuta crocuta, jaguar Panthera onca, lion Panthera leo, and tiger Panthera tigris), plus the Eurasian lynx Lynx lynx and the grey wolf Canis lupus (which are usually top predators in the areas from which data were obtained), showed greater dietary breadth and/or used a greater number of large prey species (i.e. increased generalism).
6. We suggest that dominant large carnivores encounter little competition in expanding their dietary breadth with increasing prey richness; conversely, the dietary niche of subordinate large carnivores is limited by competition with larger, dominant predators. We suggest that, over evolutionary time, resource partitioning is more important in shaping the dietary niche of smaller, inferior competitors than the niche of dominant ones.

     

Prey preferences of the spotted hyaena (Crocuta crocuta) and degree of dietary overlap with the lion (Panthera leo)

Spotted hyaenas Crocuta crocuta were once considered mere scavengers; however, detailed research revealed that they are very efficient predators. Information on what spotted hyaenas actually prefer to prey on and what they avoid is lacking, as well as the factors that influence prey selection. Data from 14 published and one unpublished study from six countries throughout the distribution of the spotted hyaena were used to determine which prey species were preferred and which were avoided using Jacobs' index. The mean of these values for each species was used as the dependent variable in multiple regression, with prey abundance and prey body mass as predictive variables. In stark contrast to the rest of Africa's large predator guild, spotted hyaenas do not preferentially prey on any species. Also surprisingly, only buffalo, giraffe and plains zebra are significantly avoided. Spotted hyaena most prefer prey within a body mass range of 56–182 kg, with a mode of 102 kg. The dietary niche breadth of the spotted hyaena is similar to that of the lion Panthera leo , and the two species have a 58.6% actual prey species overlap and a 68.8% preferred prey species overlap. These results highlight the flexible and unselective nature of spotted hyaena predation and are probably a reason for the species' success throughout its range, despite a large degree of dietary overlap with lions.     

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.     

Species richness, species range size and ecological specialisation among African primates: geographical patterns and conservation implications

The geographical distribution of species richness and species range size of African anthropoid primates (catarrhines) is investigated and related to patterns of habitat and dietary niche breadth. Catarrhine species richness is concentrated in the equatorial regions of central and west Africa; areas that are also characterised by low average species range sizes and increased ecological specificity. Species richness declines with increasing latitude north and south of the equator, while average species range size, habitat and dietary breadth increase. Relationships between species richness, species range size and niche breadth remain once latitudinal and longitudinal effects have been removed. Among areas of lowest species richness, however, there is increased variation in terms of average species range size and niche breadth, and two trends are identified. While most such areas are occupied by a few wide-ranging generalists, others are occupied by range-restricted specialist species. That conservation efforts increasingly focus on regions of high species richness may be appropriate if these regions are also characterised by species that are more restricted in both their range size and their ecological versatility, although special consideration may be required for some areas of low species richness.     

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.     

Foraging Trade‐offs between Prey Size,Delivery Rate and Prey Type: How Does Niche Breadth and Early Learning of the Foraging Niche Affect Food Delivery?

Optimal foraging theory suggests that avian parents should prefer the most energetically efficient (largest) prey items when delivering food to offspring at a central place. However, during periods of high demand, selectivity of prey may decline, leading to the delivery of smaller and/or less nutritious items. We compared foraging trade‐offs between great tits (Parus major) which had a wider feeding niche than blue tits (Cyanistes caeruleus). We also compared the foraging efficiency of cross‐fostered young, which had learned the spatial foraging niche and prey size of the foreign species, to that of control conspecifics. Mean delivery rates did not differ between control and cross‐fostered parents of either species but as delivery rates increased, prey size declined for both species and both treatment groups. However, across the range of increasing delivery rates, parents were able to increase the total biomass of prey delivered. Cross‐fostering did not alter the proportion of different prey taxa in the diet, but cross‐fostered birds shifted the size of the prey taken to that of their foster species. Consistent with their broader feeding niche, great tits, but not blue tits, incorporated more unpalatable items (flies) as delivery rates increased. Although great tits foraged less efficiently in the blue tit niche, paradoxically, blue tits seem to deliver more prey biomass when foraging in the great tit niche.     

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.     

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.     

How feeding performance and energy intake change with a small increase in the body size of the three-spined stickleback

Changes in the foraging behaviour due to variation in the body size of the three-spined stickleback Gasterosteus aculeatus were investigated. All sizes of fish had a high probability of attacking prey whenever encountered. The probability of eating the prey increased with the size of the fish, as the larger fish had larger jaws and a greater stomach capacity. Therefore, as fish increased in size there was an increase in the probability of successful prey capture. The level of satiation did not have an effect on the prey handling time, which is contrary to other studies and is probably a result of the large prey sizes. The physical size of the prey meant that the handling times were long regardless of the motivational level of the fish. The larger fish took in more energy and at a faster rate, although the time to reach satiation was similar for all fish sizes. The advantage that large fish appear to have in successfully gaining large prey is negated by their greater metabolic requirement. The changes in feeding performance induced by small increases in body size could have important consequences for intraspecific competition, habitat Use and risk of predation.     

Australian Tropical and Subtropical Rain Forest Community Assembly: Phylogeny,Functional Biogeography,and Environmental Gradients

To compare community assemblage patterns in tropical northeastern and subtropical central eastern Australia across selected gradients and scales, we tested the relationship of species traits with phylogenetic structure, and niche breadth. We considered phylogenetic relationships across current‐day species in assemblages in relation to rain forest species pool sizes, and trait values along gradients including elevation and latitude. Trait values were quantified across scales for seed size, leaf area, wood density and maximum height at maturity for 1137 species and 596 assemblages using trait gradient analysis (TGA). Local assemblages of subtropical species had narrower trait ranges, and higher niche breadth values than corresponding assemblages of tropical species. Leaf size and seed size increased at low latitudes, and community phylogenetic structure was most strongly correlated with seed traits in the subtropics, reflecting dispersal and re‐colonization processes. Elevation accounted for little of the variance in community phylogenetic structure or trait variation across local and regional scales. Stable moist forest areas retained many species from ancestral rain forest lineages across a range of temporally conserved habitats; species within assemblages were less related; and rain forest assemblages had higher functional diversity, but lower niche breadth. This suggests that on average, assemblages of species in stable areas had greater trait variation and narrower distributions. Historic and recent rain forest contraction and re‐expansion can result in recolonized areas that are dominated by species that are more related (phylogenetically) than by chance, have smaller, widely dispersed seeds, and greater niche breadth (broader distributions).     

Patterns of coexistence in sexual and asexual species of Cnemidophorus lizards

Summary The lizard genus Cnemidophorus (family Teiidae) contains sexual as well as parthenogenetic species. The theoretical two-fold fitness advantage of asexuality does not translate into any obvious distributional or numerical superiority of the parthenogenic species in the southwestern US and northern Mexico where their ranges overlap. I tested the prediction that the genetically diverse sexual species should have a higher between-individual niche width than a similar sympatric asexual species by studying the prey in stomach contents of sympatric and allopatric populations of C. tigris (sexual) and C. sonorae (asexual) in southern Arizona. The expectation proved true for niche breadths based on both prey length and prey taxa categories. The within-individual component of niche breadth was not different between species. Meaningful comparisons between species in sympatry and allopatry are confounded by the uncontrolled differences in the availability and diversity of food items between sites. Before the generality of these results can be assessed the study should be repeated in other areas where sexual and asexual species are syntopic and of similar body size.     

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.     

Predator-prey size relationships in an African large-mammal food web

1. Size relationships are central in structuring trophic linkages within food webs, leading to suggestions that the dietary niche of smaller carnivores is nested within that of larger species. However, past analyses have not taken into account the differing selection shown by carnivores for specific size ranges of prey, nor the extent to which the greater carcass mass of larger prey outweighs the greater numerical representation of smaller prey species in the predator diet. Furthermore, the top-down impact that predation has on prey abundance cannot be assessed simply in terms of the number of predator species involved. 2. Records of found carcasses and cause of death assembled over 46 years in the Kruger National Park, South Africa, corrected for under-recording of smaller species, enabled a definitive assessment of size relationships between large mammalian carnivores and their ungulate prey. Five carnivore species were considered, including lion (Panthera leo), leopard (Panthera pardus), cheetah (Acinonyx jubatus), African wild dog (Lycaon pictus) and spotted hyena (Crocuta crocuta), and 22 herbivore prey species larger than 10 kg in adult body mass. 3. These carnivores selectively favoured prey species approximately half to twice their mass, within a total prey size range from an order of magnitude below to an order of magnitude above the body mass of the predator. The three smallest carnivores, i.e. leopard, cheetah and wild dog, showed high similarity in prey species favoured. Despite overlap in prey size range, each carnivore showed a distinct dietary preference. 4. Almost all mortality was through the agency of a predator for ungulate species up to the size of a giraffe (800-1200 kg). Ungulates larger than twice the mass of the predator contributed substantially to the dietary intake of lions, despite the low proportional mortality inflicted by predation on these species. Only for megaherbivores substantially exceeding 1000 kg in adult body mass did predation become a negligible cause of mortality. 5. Hence, the relative size of predators and prey had a pervasive structuring influence on biomass fluxes within this large-mammal food web. Nevertheless, the large carnivore assemblage was dominated overwhelmingly by the largest predator, which contributed the major share of animals killed across a wide size range.     

Niche overlap of two sympatric-nesting hawks Accipiter spp. in the New Jersey-New York Highlands

The food and habitat niches of two nesting species of hawks Accipiter spp were studied in an extensively forested area of the Eastern Deciduous Forest Biome Nesting habitat was quantitated at 19 Cooper s hawk A cooperu nests and 16 northern goshawk A gentilis nests There was no significant trend for Cooper's hawks to nest in less mature forests than northern goshawks as reported previously for western North America Forest habitats did not differ markedly except that shrub cover was greater at Cooper's hawk nest sites, which were also on flatter terrain and closer to roads, forest openings, and human habitation However, these few differences resulted m reducing habitat-niche overlap considerably (0 538), as was calculated using principal components analysis Mean prey weight was significantly larger for the northern goshawk which follows its 2 2-fold body weight advantage over Cooper's hawk Although bird prey was of primary importance to both Accipiter, goshawks took twice the proportion of mammals compared to their smaller congener Food-niche overlap was lowest by prey species overlap (0 470), followed by prey size class overlap (0 529), and highest by vertical foraging zone overlap (0 816) The Cooper's hawk showed the greatest niche breadth for both food and habitat niches indicating it as more of a generalist Overall, niche complementarity of food and habitat dimensions resulted in niche overdispersion along food and habitat dimensions with a total niche overlap (0 504) that was below the competition threshold These results suggested that competition (past and current) was responsible for segregating niches