Food webs robustness to biodiversity loss: the roles of connectance, expansibility and degree distribution

We analyse the robustness of food webs against species loss by considering the influence of several structural factors of the networks, such as connectance, degree distribution and expansibility. The last concept refers to the absence of structural bottlenecks in the food web, whose removal separate the network into large isolate clusters. In theory networks with identical connectance can display different expansibility characteristics. Using the spectral scaling method we studied 17 food networks and classified them as good expansion (GE) and not-GE networks. The combination of GE properties and degree distribution of species permitted the classification of food webs into six different classes. These classes characterize the differences in robustness of food webs to species loss. While the webs having uniform degree distributions and displaying GE properties are the most robust to species loss, the presence of bottlenecks and skewed distribution of the number of links per species make food webs very vulnerable to primary removal of species.     

Biogeographic variation of food habits and body size of the America puma

Summary The puma (Felis concolor) has the most extensive range of any terrestrial mammal in the Western Hemisphere, covering over 100° latitude. Food habits of different puma subspecies vary with latitude. Subspecies from temperate habitats generally eat larger prey and specialize on a smaller number of prey taxa, whereas, in tropical habitats, they prey on smaller, more varied prey. In North America, ungulates (primarily deer) represented 68% of the puma's diet by frequency of occurrence. Mean weight of vertebrate prey (MWVP) was positively correlated (r=0.875) with puma body weight and inversely correlated (r=-0.836) with food niche breadth in all America. In general, MWVP was lower in areas closer to the Equator. Patterns of puma prey selection are probably influenced by prey availability and vulnerability, habitat characteristics, and potential competition from the jaguar (Panthera onca).     

Food webs: experts consuming families of experts

Food webs of habitats as diverse as lakes or desert valleys are known to exhibit common "food-web patterns", but the detailed mechanisms generating these structures have remained unclear. By employing a stochastic, dynamical model, we show that many aspects of the structure of predatory food webs can be understood as the traces of an evolutionary history where newly evolving species avoid direct competition with their relatives. The tendency to avoid sharing natural enemies (apparent competition) with related species is considerably weaker. Thus, "experts consuming families of experts" can be identified as the main underlying food-web pattern. We report the results of a systematic, quantitative model validation showing that the model is surprisingly accurate.     

食物浓度对角突臂尾轮虫种群增长、个体大小和卵大小的影响

以蛋白核小球藻为食物 ,应用群体累积培养法研究了食物浓度对角突臂尾轮虫种群增长、个体大小和卵大小等的影响 .结果表明 ,食物浓度对角突臂尾轮虫种群增长率、个体大小和卵大小均有极显著影响 .其中 ,轮虫种群增长率与食物浓度间呈曲线相关 ;当食物浓度为 8.2 4 5 3× 10 6cells·ml-1时 ,种群增长率达最大值 0 .6 0 85d-1;在所研究的食物浓度范围内 ,轮虫个体具有随食物浓度的增大而增大的趋势 ,轮虫卵体积在中等食物浓度范围 (6 .0× 10 6～ 9.0× 10 6cells·ml-1)时较大     

Reproductive strategies of primates: The influence of body size and diet on litter size

The frequency of multiple births, life history parameters, body size, and diet characteristics were obtained from the literature for 70 primate species. The general pattern within the primate order is to have single infant litters, yet multiple births regularly occur in a number of species in specific phylogenetic groups. Primates which have large litters tend to be small, have short gestation periods and give birth to small infants, which are weaned quickly, and mature rapidly. Species in which multiple births are common also have short interbirth intervals and in the Callitrichidae have males which exhibit paternal care. In addition, they are commonly insectivorous. Although it is difficult to isolate the effects of diet on litter size, independent of body size, analyses suggest that after the influence of body size is statistically removed, as the proportion of insects in the diet increases, animals have larger litters. We suggest that by adopting a mixed diet of insects and fruit primates may be able to ensure access to a seasonally stable food resource that is not greatly restricted by the presence of toxins. This diet would allow a relatively high metabolism and facilitate large litters.     

The effects of energy input,immigration and habitat size on food web structure: a microcosm experiment

It has been hypothesised that larger habitats should support more complex food webs. We consider three mechanisms which could lead to this pattern. These are increased immigration rates, increased total productivity and spatial effects on the persistence of unstable interactions. Experiments designed to discriminate between these mechanisms were carried out in laboratory aquatic microcosm communities of protista and bacteria, by independently manipulating habitat size, total productivity and immigration rate. Larger habitats supported more complex food webs, with more species, more links per species and longer maximum and mean food chains, even in the absence of differences in total energy input. Increased immigration rate resulted in more complex food webs, but habitats with higher energy input per unit area supported less complex food webs. We conclude that spatial effects on the persistence of unstable interactions, and variation in immigration rates, are plausible mechanisms by which habitat size could affect food web structure. Variation in total productivity with habitat area seems a less likely explanation for variation in food web structure.     

Contrasting patterns of body size for Daphnia species that segregate by habitat

Summary We investigate how body size of two coexisting Daphnia species varies among 7 lakes that represent a gradient of predation risk. The two species segregate vertically in stratified lakes; D. galeata mendotae is typically smaller and more eplimnetic than D. pulicaria. The extent of vertical habitat partitioning, however, varies seasonally within and among lakes in apparent response to predation intensity by epilimnetic planktivorous fishes. Daphnia pulicaria uses the epilimnion at low levels of fish predation but is restricted to the hypolimnion under high fish predation, whereas D. galaeta mendotae always utilizes the epilimnion. The species display contrasting patterns of genetic variation in neonate size and size at maturity. D. pulicaria is larger in lakes with higher fish and Chaoborus densities whereas D. galeata mendotae is smaller. This contrast in body size in lakes with high predation is associated with greater habitat segregation in those lakes. In lakes with low predation risk, the two species are similar in body size at birth and maturity.Authorship order alphabetical     

On the importance of body size in the colonisation of ephemeral resource patches by vagile consumers

A study on the colonisation of leaf detritus patches by vagile macroinvertebrates in a brackish lagoon is presented in the framework of a conceptual model where a body size-related constraint on patch use behaviour is explicitly considered. Abundance patterns of dominant macroinvertebrate taxa were characterised by short-term, non-random fluctuations, showing significant site-dependent variations. Yet, a site-independent covariation was observed between patterns’ fractal dimension and the average body mass of each taxon, indicating that, while the temporal scales characterising the colonisation patterns may be highly species-specific, cross-species generalisations are possible based on body size. The generality of these results was supported by literature data on temporal patterns of carcass colonisation by bathyal fish. The importance of size-related mechanisms in regulating the aggregation of vagile consumers on resource patches and, ultimately, their coexistence at both an inter- and intra-specific level, is discussed.     

On the role of body size in a tri-trophic metapopulation model

 A particular tri-trophic (resource, prey, predator) metapopulation model with dispersal of preys and predators is considered in this paper. The analysis is carried out numerically, by finding the bifurcations of the equilibria and of the limit cycles with respect to prey and predator body sizes. Two routes to chaos are identified. One is characterized by an intriguing cascade of flip and tangent bifurcations of limit cycles, while the other corresponds to the crisis of a strange attractor. The results are summarized by partitioning the space of body sizes in eight subregions, each one of which is associated to a different asymptotic behavior of the system. Emphasis is put on the possibility of having different modes of coexistence (stationary, cyclic, and chaotic) and/or extinction of the predator population. Received 1 August 1995; received in revised form 8 January     

Fish predation on Eudiaptomus gracilis in relation to clutch size,body size,and sex: a field experiment

An enclosure experiment was performed to test for direct predationeffects on fecundity and adult body size of the copepod Eudiaptomus gracilis in the field. By introducing a high densityof fish (15 underyearling roach, Rutilus rutilus, per 350litre enclosure) and documenting the short-term effects on traitsin a rapidly decreasing prey population, responses to changes inthe phytoplankton community were minimized. After 68 hours ofpredation, clutch size and frequency of females carrying eggs weresignificantly lower in fish enclosures. Female density was moreaffected than male density. Predation selected against large bodysize in both sexes but less so in females, leading to an increasedsexual size dimorphism. The results agree with predictions based onprey selectivity in fish. Predation risk should increase withclutch size and body size since these traits increase theconspicuousness of prey. The size of the highly visible egg-clutchmay be more important than body size. Female body size wasuncorrelated to clutch size, which may explain the weaker sizeeffect among females and the changed sexual sizedimorphism.     

The relationship between ecological segregation and sexual body size dimorphism in large herbivores

Ecological segregation (sexual differences in diet or habitat use) in large herbivores has been intimately linked to sexual body size dimorphism, and may affect both performance and survival of the sexes. However, no one has tested comparatively whether segregation occurs at a higher frequency among more dimorphic species. To test this comparatively, data on sex-specific diet, habitat use and body size of 40 species of large herbivores were extracted from the literature. The frequency of ecological segregation was higher among more dimorphic herbivores; however, this was only significant for browsers. This provides the first evidence that segregation is more common among more dimorphic species. The comparative evidence supported the nutritional-needs hypothesis over the incisor breadth hypothesis, as there was no difference in frequency of segregation between seasons with high and low resource levels, and since segregation was also evident among browsers. Whether the absence of a correlation between ecological segregation and level of sexual body size dimorphism for intermediate feeders and grazers is due to biological differences relative to browsers or to the fact that the monomorphic species included in the analysis were all browsers is discussed. Received: 18 August 1999 / Accepted: 31 January 2000     

Niche differentiation depends on body size in a cichlid fish: a model system of a community structured according to size regularities

1. Communities of different species are often structured according to niche differentiation associated with competitive interactions. We show that similar principles may apply on an ecological time-scale when individuals of a species having a wide size variation compete for resources, using the Lake Tanganyika cichlid Lobochilotes labiatus (5-30 cm). This species has a mouth especially adapted to suck up invertebrates from rock crevices. 2. Individuals defended feeding territories against similar-sized conspecifics, but not against different-sized ones. Thus, territories of similar-sized fish rarely overlapped, but up to a total of seven individuals (of seven size-ranks) had broadly overlapping territories with dissimilar-sized individuals. Comparison with expectation from the null model demonstrated clearly that observed size ratios between adjacent size rank were determined non-randomly regardless of sexual combinations. 3. Larger individuals took larger prey types of larger average size, but more importantly used wider rock crevices from which to suck food than smaller individuals. We calculated pairwise values of Schoener's index of diet overlap C(d) and the values of Levin's index of diet breadth B(d) (prey type and prey size) and the same for the width of the rock crevices used for foraging (C(r) and B(r)). C(d) remained high among all combinations of the seven ranks. In contrast, C(r) declined strongly in combinations of adjacent ranks (to 0.27), and was low or zero among further different size ranks. This shows that fish with overlapping territories divided the food resources largely through foraging site partitioning. Accordingly, B(d) did not depend on the size difference to the nearest two coinhabiting fish, whereas B(r) did. 4. We conclude that this L. labiatus community is structured non-randomly: body size-dependent effects on foraging site usage result in competition with, and territorial exclusion of, similar-sized individuals, but not of dissimilar-sized individuals that were accepted as coinhabitants. Accordingly, mean body size ratios (large/small) between two adjacent ranks were consistently approximately 1.28 [standard deviation (SD) = 0.07, n = 104], while approximately 1.34 from the null model (SD = 0.34, n = 10 400 simulations). We discuss our results as an example of Hutchinson's rule, applied originally to size ratios of different species.     

Resolving the roles of body size and species identity in driving functional diversity

Efforts to characterize food webs have generated two influential approaches that reduce the complexity of natural communities. The traditional approach groups individuals based on their species identity, while recently developed approaches group individuals based on their body size. While each approach has provided important insights, they have largely been used in parallel in different systems. Consequently, it remains unclear how body size and species identity interact, hampering our ability to develop a more holistic framework that integrates both approaches. We address this conceptual gap by developing a framework which describes how both approaches are related to each other, revealing that both approaches share common but untested assumptions about how variation across size classes or species influences differences in ecological interactions among consumers. Using freshwater mesocosms with dragonfly larvae as predators, we then experimentally demonstrate that while body size strongly determined how predators affected communities, these size effects were species specific and frequently nonlinear, violating a key assumption underlying both size- and species-based approaches. Consequently, neither purely species- nor size-based approaches were adequate to predict functional differences among predators. Instead, functional differences emerged from the synergistic effects of body size and species identity. This clearly demonstrates the need to integrate size- and species-based approaches to predict functional diversity within communities.     

Emergence of complexity in evolving niche-model food webs

We have analysed mechanisms that promote the emergence of complex structures in evolving model food webs. The niche model is used to determine predator-prey relationships. Complexity is measured by species richness as well as trophic level structure and link density. Adaptive dynamics that allow predators to concentrate on the prey species they are best adapted to lead to a strong increase in species number but have only a small effect on the number and relative occupancy of trophic levels. The density of active links also remains small but a high number of potential links allows the network to adjust to changes in the species composition (emergence and extinction of species). Incorporating effects of body size on individual metabolism leads to a more complex trophic level structure: both the maximum and the average trophic level increase. So does the density of active links. Taking body size effects into consideration does not have a measurable influence on species richness. If species are allowed to adjust their foraging behaviour, the complexity of the evolving networks can also be influenced by the size of the external resources. The larger the resources, the larger and more complex is the food web it can sustain. Body size effects and increasing resources do not change size and the simple structure of the evolving networks if adaptive foraging is prohibited. This leads to the conclusion that in the framework of the niche model adaptive foraging is a necessary but not sufficient condition for the emergence of complex networks. It is found that despite the stabilising effect of foraging adaptation the system displays elements of self-organised critical behaviour.     

Body size and ecological diversification in a sister species pair of triplefin fishes

The effect of body size on spatial resource competition and reproductive isolation was examined in a sister species pair of subtidal triplefin fishes (F. Tripterygiidae) in New Zealand. Ruanoho decemdigitatus and Ruanoho whero have overlapping sympatric distributions and differ in body size, attaining a total length of 12 cm and 9 cm, respectively. R. decemdigitatus was most commonly found in sheltered areas shallower than 5 m, while R. whero was frequently found in sheltered to moderately exposed areas down to 20 m. In sites where the species co-occurred, R. whero was less associated with rock substratum. The effect of body size on substratum use was investigated using laboratory trials based on the field data to test habitat preference and competitive ability in a common setting. Reproductive behaviour was assessed in courtship, mate choice and hybridisation trials. Both species exhibited similar habitat preferences, but large R. decemdigitatus were dominant in inter- and intraspecific contests for the preferred rock habitat, while small R. whero were displaced into less preferred habitats. Courtship behaviour in R. whero was a subset of that displayed by R. decemdigitatus, while no mating behaviour was observed in heterospecific trials. Female R. whero showed a strong preference for smaller males, while female R. decemdigitatus had no preference for male size. Results suggest that body size differences in the Ruanoho pair are consistent with female choice for smaller males in R. whero and competition for habitat in both species. Body size in the Ruanoho species appears to be influenced by conflicting selection pressures that may differ between the species.     

Relating middle-ear acoustic performance to body size in the cat family: measurements and models

Is the acoustic performance of the mammalian middle ear dependent on body size? We focus on the cat family, because of its qualitatively uniform (and distinctive) middle-ear structure, large size range, and the extensive data available from domestic cats which provide a framework for relating middle-ear acoustics to structure. We report measurements of acoustic admittance in 17 live adult ears of 11 exotic species, ranging in size from sand cat (3?kg) to tiger (180?kg). For low frequencies, the middle-ear response is compliant for all species and generally increases with size. The compliance of the middle-ear air space increases with size, but the compliance of the tympanic membrane and ossicular chain is not correlated with size. Structure-based rules are developed to represent some features of middle-ear performance: (1) low-frequency sensitivity increases with size; and (2) the frequency of a prominent notch in admittance decreases with size. Although some species deviate from the rules, the data generally support the idea that in larger felids the middle-ear response is shifted to lower frequencies. Thus, in the cat family, body size partly describes variations in auditory features. More speculatively, ethological pressures which might influence hearing performance are discussed.     

Joint evolution of predator body size and prey-size preference

We studied the joint evolution of predator body size and prey-size preference based on dynamic energy budget theory. The predators’ demography and their functional response are based on general eco-physiological principles involving the size of both predator and prey. While our model can account for qualitatively different predator types by adjusting parameter values, we mainly focused on ‘true’ predators that kill their prey. The resulting model explains various empirical observations, such as the triangular distribution of predator–prey size combinations, the island rule, and the difference in predator–prey size ratios between filter feeders and raptorial feeders. The model also reveals key factors for the evolution of predator–prey size ratios. Capture mechanisms turned out to have a large effect on this ratio, while prey-size availability and competition for resources only help explain variation in predator size, not variation in predator–prey size ratio. Predation among predators is identified as an important factor for deviations from the optimal predator–prey size ratio.     

Food web structure and interaction strength pave the way for vulnerability to extinction

This paper focuses on how food web structure and interactions among species affects the vulnerability, due to environmental variability, to extinction of species at different positions in model food webs. Vulnerability is here not measured by a traditional extinction threshold but is instead inspired by the IUCN criteria for endangered species: an observed rapid decline in population abundance. Using model webs influenced by stochasticity with zero autocorrelation, we investigate the ecological determinants of species vulnerability, i.e. the trophic interactions between species and food web structure and how these interact with the risk of sudden drops in abundance of species. We find that (i) producers fulfil the criterion of vulnerable species more frequently than other species, (ii) food web structure is related to vulnerability, and (iii) the vulnerability of species is greater when involved in a strong trophic interaction than when not. We note that our result on the relationship between extinction risk and trophic position of species contradict previous suggestions and argue that the main reason for the discrepancy probably is due to the fact that we study the vulnerability to environmental stochasticity and not extinction risk due to overexploitation, habitat destruction or interactions with introduced species. Thus, we suggest that the vulnerability of species to environmental stochasticity may be differently related to trophic position than the vulnerability of species to other factors. Earlier research on species extinctions has looked for intrinsic traits of species that correlate with increased vulnerability to extinction. However, to fully understand the extinction process we must also consider that species interactions may affect vulnerability and that not all extinctions are the result of long, gradual reductions in species abundances. Under environmental stochasticity (which importance frequently is assumed to increase as a result of climate change) and direct and indirect interactions with other species some extinctions may occur rapidly and apparently unexpectedly. To identify the first declines of population abundances that may escalate and lead to extinctions as early as possible, we need to recognize which species are at greatest risk of entering such dangerous routes and under what circumstances. This new perspective may contribute to our understanding of the processes leading to extinction of populations and eventually species. This is especially urgent in the light of the current biodiversity crisis where a large fraction of the world's biodiversity is threatened.     

Adaptive significance of small body size under poor socio-economic conditions in Southern Peru

The relationship of variations in parental body size to offspring survival has been studied in a population of poor socio-economic conditions (“Barriada”) in the southern highland of Peru. Parents of small body size, especially mothers, had significantly greater per cent offspring survival than parents with larger body size. In other words, the offspring survival effectiveness of subjects of small body size was greater than that of subjects of large body size. It is postulated that the greater offspring survival effectiveness associated with small parental body size may reflect possible adaptive responses to poor socio-economic conditions.