Group formation stabilizes predator-prey dynamics

Theoretical ecology is largely founded on the principle of mass action, in which uncoordinated populations of predators and prey move in a random and well-mixed fashion across a featureless landscape. The conceptual core of this body of theory is the functional response, predicting the rate of prey consumption by individual predators as a function of predator and/or prey densities. This assumption is seriously violated in many ecosystems in which predators and/or prey form social groups. Here we develop a new set of group-dependent functional responses to consider the ecological implications of sociality and apply the model to the Serengeti ecosystem. All of the prey species typically captured by Serengeti lions (Panthera leo) are gregarious, exhibiting nonlinear relationships between prey-group density and population density. The observed patterns of group formation profoundly reduce food intake rates below the levels expected under random mixing, having as strong an impact on intake rates as the seasonal migratory behaviour of the herbivores. A dynamical system model parameterized for the Serengeti ecosystem (using wildebeest (Connochaetes taurinus) as a well-studied example) shows that grouping strongly stabilizes interactions between lions and wildebeest. Our results suggest that social groups rather than individuals are the basic building blocks around which predator-prey interactions should be modelled and that group formation may provide the underlying stability of many ecosystems.     

Predator diversity dampens trophic cascades

Food web complexity is thought to weaken the strength of terrestrial trophic cascades in which strong impacts of natural enemies on herbivores cascade to influence primary production indirectly. Predator diversity can enhance food web complexity because predators may feed on each other and on shared prey. In such cases, theory suggests that the impact of predation on herbivores relaxes and cascading effects on basal resources are dampened. Despite this view, no empirical studies have explicitly investigated the role of predator diversity in mediating primary productivity in a natural terrestrial system. Here we compare, in a coastal marsh community, impacts of arthropod predators on herbivores and plant productivity between a simple food web with a single predator species and a complex food web with a diverse predator assemblage. We show that enhancing predator diversity dampens enemy effects on herbivores and weakens trophic cascades. Consequently, changes in diversity at higher trophic levels can significantly alter ecosystem function in natural systems.     

Allometric degree distributions facilitate food-web stability

In natural ecosystems, species are linked by feeding interactions that determine energy fluxes and create complex food webs. The stability of these food webs enables many species to coexist and to form diverse ecosystems. Recent theory finds predator-prey body-mass ratios to be critically important for food-web stability. However, the mechanisms responsible for this stability are unclear. Here we use a bioenergetic consumer-resource model to explore how and why only particular predator-prey body-mass ratios promote stability in tri-trophic (three-species) food chains. We find that this 'persistence domain' of ratios is constrained by bottom-up energy availability when predators are much smaller than their prey and by enrichment-driven dynamics when predators are much larger. We also find that 97% of the tri-trophic food chains across five natural food webs exhibit body-mass ratios within the predicted persistence domain. Further analyses of randomly rewired food webs show that body mass and allometric degree distributions in natural food webs mediate this consistency. The allometric degree distributions hold that the diversity of species' predators and prey decreases and increases, respectively, with increasing species' body masses. Our results demonstrate how simple relationships between species' body masses and feeding interactions may promote the stability of complex food webs.     

Mineral nutrition and spatial concentrations of African ungulates

Africa's abundant large herbivores are very heterogeneously distributed, both geographically and regionally. Within a region, some localities contain dense animal concentrations although areas nearby may be virtually unoccupied. Mixed-species herds are a conspicuous feature of areas where animals concentrate. The prevailing explanations of local distributional concentrations are (1) that different herbivore species facilitate each other's foraging, and (2) that animals are protected from predation by both intraspecific and interspecific association. If facilitation of grazing were an overriding factor, mixed species herds should move extensively with localized rain showers to obtain the greatest forage yield. If predation were the major factor influencing animal densities and distributions, rapid, unpredictable spatial movements would further reduce predation. But because resident, non-migratory species tend to occupy home ranges that are stable over time, neither of these hypotheses is totally compelling. Because tropical forages are of lower quality than temperate ones and are often chronically deficient in mineral elements, I tested the hypothesis that areas where animals concentrate are localities supporting forages of higher mineral content. I report here that the mineral content of foods is an important determinant of the spatial distributions of animals within the Serengeti National Park, Tanzania. Based on ecological critieria, magnesium, sodium and phosphorus appear particularly important.     

A comparative analysis of the evolution of imperfect mimicry

Although exceptional examples of adaptation are frequently celebrated, some outcomes of natural selection seem far from perfect. For example, many hoverflies (Diptera: Syrphidae) are harmless (Batesian) mimics of stinging Hymenoptera. However, although some hoverfly species are considered excellent mimics, other species bear only a superficial resemblance to their models and it is unclear why this is so. To evaluate hypotheses that have been put forward to explain interspecific variation in the mimetic fidelity of Palearctic Syrphidae we use a comparative approach. We show that the most plausible explanation is that predators impose less selection for mimetic fidelity on smaller hoverfly species because they are less profitable prey items. In particular, our findings, in combination with previous results, allow us to reject several key hypotheses for imperfect mimicry: first, human ratings of mimetic fidelity are positively correlated with both morphometric measures and avian rankings, indicating that variation in mimetic fidelity is not simply an illusion based on human perception; second, no species of syrphid maps out in multidimensional space as being intermediate in appearance between several different hymenopteran model species, as the multimodel hypothesis requires; and third, we find no evidence for a negative relationship between mimetic fidelity and abundance, which calls into question the kin-selection hypothesis. By contrast, a strong positive relationship between mimetic fidelity and body size supports the relaxed-selection hypothesis, suggesting that reduced predation pressure on less profitable prey species limits the selection for mimetic perfection.     

Predators increase the risk of catastrophic extinction of prey populations.

There has been considerable research on both top-down effects and on disturbances in ecological communities; however, the interaction between the two, when the disturbance is catastrophic, has rarely been examined. Predators may increase the probability of prey extinction resulting from a catastrophic disturbance both by reducing prey population size and by changing ecological traits of prey individuals such as habitat characteristics in a way that increases the vulnerability of prey species to extinction. We show that a major hurricane in the Bahamas led to the extinction of lizard populations on most islands onto which a predator had been experimentally introduced, whereas no populations became extinct on control islands. Before the hurricane, the predator had reduced prey populations to about half of those on control islands. Two months after the hurricane, we found only recently hatched individuals--apparently lizards survived the inundating storm surge only as eggs. On predator-introduction islands, those hatchling populations were a smaller fraction of pre-hurricane populations than on control islands. Egg survival allowed rapid recovery of prey populations to pre-hurricane levels on all control islands but on only a third of predator-introduction islands--the other two-thirds lost their prey populations. Thus climatic disturbance compounded by predation brought prey populations to extinction.     

Snake circumvents constraints on prey size

For animals who are unable to take bites out of their food, the size of the food item that can be consumed is constrained by the maximal size of the mouth opening (gape)--snakes are an example of gape-limited predators and they usually swallow their prey whole. Here we describe unique feeding behaviours in two closely related species of snake, which circumvent their gape limitation by removing and consuming pieces from newly moulted crabs that are too large to be swallowed intact. This evolutionary innovation is surprising, as the needle-like teeth and highly mobile bones that facilitate the capture and engulfment of large, whole prey by snakes are ill-suited both to cutting and to generating large bite forces.     

The effects of competition and predation on diversification in a model adaptive radiation

Much of life's diversity is thought to have arisen through successive rounds of adaptive radiation-the rapid diversification of a lineage into a range of ecologically and phenotypically distinct species. Both resource competition and predation have been suggested as mechanisms driving this process, although the former is better studied than the latter. Here we show experimentally how predation by a protist, Tetrahymena thermophila, affects diversification in a model adaptive radiation of the bacterial prey, Pseudomonas fluorescens. We estimate the frequency-dependent fitness functions of competing niche-specialist prey in the presence and absence of predation, and use these to test hypotheses about the extent (measured as the number of new genotypes) and rate of diversification. Competition and predation independently generated diversifying selection that we show is capable of driving prey diversification to similar extents but at different rates, diversification being markedly delayed in the presence of predators. The cause of this delay stems from weaker diversifying selection due to the reduction in prey density caused by predation. Our results suggest that predation may play an under-appreciated role in driving adaptive radiations.     

捕食者追捕与食饵逃逸对共位捕食系统中物种共存的影响

基于元胞自动机模型,探讨捕食者追捕和食饵逃逸两种行为对共位捕食系统中物种共存动态的影响.结果表明:捕食者追捕能够扩大物种共存区域,但食饵逃逸能否提高共位捕食系统中物种的共存取决于两物种竞争与捕食能力的相互权衡关系.在捕食者占绝对优势以及竞争与捕食之间存在弱权衡关系时,食饵逃逸能够促进两物种的共存;而在食饵占有绝对优势以及两物种间强权衡关系时,食饵逃逸则使得物种共存变得更加困难.另外,模拟结果显示,捕食者追捕与食饵逃逸对物种共存与种群密度产生不对称性效应:捕食者追捕对系统产生的影响较食饵逃逸行为更为显著.     

Energetic constraints on the diet of terrestrial carnivores

Species in the mammalian order Carnivora exhibit a huge diversity of life histories with body sizes spanning more than three orders of magnitude. Despite this diversity, most terrestrial carnivores can be classified as either feeding on invertebrates and small vertebrates or on large vertebrates. Small carnivores feed predominantly on invertebrates probably because they are a superabundant resource (sometimes 90% of animal biomass); however, intake rates of invertebrate feeders are low, about one tenth of those of vertebrate feeders. Although small carnivores can subsist on this diet because of low absolute energy requirements, invertebrate feeding appears to be unsustainable for larger carnivores. Here we show, by reviewing the most common live prey in carnivore diets, that there is a striking transition from feeding on small prey (less than half of predator mass) to large prey (near predator mass), occurring at predator masses of 21.5-25 kg. We test the hypothesis that this dichotomy is the consequence of mass-related energetic requirements and we determine the predicted maximum mass that an invertebrate diet can sustain. Using a simple energetic model and known invertebrate intake rates, we predict a maximum sustainable mass of 21.5 kg, which matches the point where predators shift from small to large prey.     

The evolution of müllerian mimicry in multispecies communities

Prey species that are unprofitable to attack often share conspicuous colours and patterns with other coexisting defended species. This phenomenon, termed müllerian mimicry, has long been explained as a consequence of selection on defended prey to adopt a common way of advertising their unprofitability. However, studies using two unpalatable prey types have not always supported this theory. Here we show, using a system of humans hunting for computer-generated prey, that predators do not always generate strong selection for mimicry when there are two unprofitable prey types. By contrast, we demonstrate that when predators are faced with a range of different prey species, selection on unprofitable prey to resemble one another can be intense. Here the primary selective force is not one in which predators evaluate the profitabilities of distinct prey types independently, but one in which predators learn better to avoid unprofitable phenotypes that share traits distinguishing them from profitable prey. This need to simplify decision making readily facilitates the spread of imperfect mimetic forms from rarity, and suggests that müllerian mimicry is more likely to arise in multispecies communities.     

Visual predators select for crypticity and polymorphism in virtual prey

Cryptically coloured animals commonly occur in several distinct pattern variants. Such phenotypic diversity may be promoted by frequency-dependent predation, in which more abundant variants are attacked disproportionately often, but the hypothesis has never been explicitly tested. Here we report the first controlled experiment on the effects of visual predators on prey crypticity and phenotypic variance, in which blue jays (Cyanocitta cristata) searched for digital moths on computer monitors. Moth phenotypes evolved via a genetic algorithm in which individuals detected by the jays were much less likely to reproduce. Jays often failed to detect atypical cryptic moths, confirming frequency-dependent selection and suggesting the use of searching images, which enhance the detection of common prey. Over successive generations, the moths evolved to become significantly harder to detect, and they showed significantly greater phenotypic variance than non-selected or frequency-independent selected controls.     

The role of parasites in sympatric and allopatric host diversification

Exploiters (parasites and predators) are thought to play a significant role in diversification, and ultimately speciation, of their hosts or prey. Exploiters may drive sympatric (within-population) diversification if there are a variety of exploiter-resistance strategies or fitness costs associated with exploiter resistance. Exploiters may also drive allopatric (between-population) diversification by creating different selection pressures and increasing the rate of random divergence. We examined the effect of a virulent viral parasite (phage) on the diversification of the bacterium Pseudomonas fluorescens in spatially structured microcosms. Here we show that in the absence of phages, bacteria rapidly diversified into spatial niche specialists with similar patterns of diversity across replicate populations. In the presence of phages, sympatric diversity was greatly reduced, as a result of phage-imposed reductions in host density decreasing competition for resources. In contrast, allopatric diversity was greatly increased as a result of phage-imposed selection for resistance, which caused populations to follow divergent evolutionary trajectories. These results show that exploiters can drive diversification between populations, but may inhibit diversification within populations by opposing diversifying selection that arises from resource competition.     

Multiple benefits of gregariousness cover detectability costs in aposematic aggregations

Understanding the early evolution of aposematic (warning) coloration has been a challenge for scientists, as a new conspicuous morph in a population of cryptic insects would have a high predation risk and would probably die out before local predators learnt to avoid it. Fisher presented the idea of aggregation benefit through the survival of related individuals; however, his theory has been strongly debated as the mechanisms that favour grouping have never been explored experimentally with the incorporation of detectability costs. Here we create a comprehensive 'novel world' experiment with the great tit (Parus major) as a predator to explore simultaneously the predation-related benefits and costs for aposematic aggregated prey, manipulating both group size and signal strength. Our results show that grouping would have been highly beneficial for the first aposematic prey individuals surrounded by naive predators, because (1) detectability risk increased only asymptotically with group size; (2) additional detectability costs due to conspicuous signals were marginal in groups; (3) even naive predators deserted the group after detecting unpalatability (dilution effect); and (4) avoidance learning of signal was faster in groups. None of these mechanisms require kin selection.     

依赖比率的捕食被捕食系统的全局分析:捕食与被捕食者均有密度制约的情况

近年来依赖比率的捕食与被捕食模型已引起了生态学者和生物数学研究者的密切注意，这是因为它们比传统的模型更加接近实际情形．本文研究了依赖比率的捕食与被捕食系统的全局定性性态，这里捕食与被捕食者均为密度制约的．     

Phylogeny and life habits of Early Arthropods-Predation in the Early Cambrian Sea

We investigated two new arthropods from the Maotianshan-Shale fauna of southern China in the course of our research on life strategies, particularly predation, in Early Cambrian marine macrofaunal biota. One form clearly belongs to the so-called "great-appendage" arthropods, animals that were, most likely, active predators catching prey with their first pair of large, specialized frontoventral appendages. Based on this, we hypothesize that the new species and many others, if not all, of the "great-appendage" arthropods were derivatives of the chelicerate stem lineage and not forms having branched off at different nodes along the evolutionary lineage of the Arthropoda. Rather, we consider the "great-appendage" arthropods as belonging to a monophyletic clade, which modified autapomorphically their first pair of appendages (antennae in general arthropod terminology) into raptorial organs for food capture. The second new form resembles another Maotianshan-Shale arthropod, Fuxianhuia protensa, in sharing a head made of only two separate segments, a small segment bearing oval eyes laterally, and another bearing a large tergite, which forms a wide shield freely overhanging the subsequent narrow trunk segments. This segment bears a single pair of rather short anteriorly directed uniramous appendages, considered as the "still" limb-shaped antennae. Particularly the evolutionary status of head and limbs of these two forms suggests that both are representatives of the early part of the stem lineage toward the crown-group of Arthropoda, the Euarthropoda. These forms appear rather unspecialized, but may have been but simple predators. This adds to our hypothesis that predation was a common, if not dominant feeding strategy in the Cambrian, at least for arthropods.     

越冬长耳鸮(Asio otus)食物分析——以上海江苏两地样本为例

2010—2012年、2014—2015年,采用食团内容物分析法分别对上海崇明、江苏盐城越冬长耳鸮的食物组成进行分析.两个越冬地3年时间内共收集完整食团717份,辨认出7种804只动物个体,其中鼠类、麝鼩等小型哺乳动物4种,占总数的93.7%,雀形目鸟类3种,及少量无法鉴定到种的鸟类.结果表明,两地越冬的长耳鸮对黑线姬鼠(Apodemus agrarius)的捕食量最大,捕食数量占食物总数的89.2%;对黑线姬鼠成年Ⅰ组的捕食比例最高,为40.2%;对黑线姬鼠各年龄组的捕食量与其当地种群数量变化基本一致.由此推测,越冬长耳鸮捕食的主要对象与周围环境中可获得的食物来源有关.     

Influence of long-period pesticide force on genetic polymorphism of wolf spider Pardosa pseudoannulata (Lycosidae: Araneae)

Wolf spiders are predators in large quantities in the fields. How wolf spiders keep their dominant species status under long-period pesticide force? To answer this question, eight geographical populations of Pardosa pseudoannulata were used as materials to test the influence of geographical habitats on their genomic DNA polymorphism. The RAPD pattern showed polymorphic variations among and within different populations. Total 84 bands amplified by 10 random primers, of which 62 (73.81%) are polymorphic, were generated from 55 individuals of eight geographical populations. Meanwhile, Shannon’s index (Ho= 0.5177) showed a rich genetic diversity of P. pseudoannulata, and most of the genetic variation (64.24%) was found within populations. Multiple regression analysis suggested that it is the climatic variation (such as annual average temperature etc.) that results in adaptive eco-geographic differentiation, and it is the long-period pesticide force that speeds up the genetic differentiation of P. pseudoannulata which changed the genetic diversity of the population.     

Post-spawning egg care by a squid

Gonatus onyx is one of the most abundant cephalopods in the Pacific and Atlantic Oceans and is an important prey species for a variety of vertebrate predators, but a full understanding of its life history has been hampered because spawning occurs at great depths, where observation is difficult. Here we describe post-spawning egg care, or brooding, in this deep-sea squid. Our finding is unexpected because this behaviour differs from the reproductive habits of all other known squid species.     

Ecology: is speciation driven by species diversity?

Emerson and Kolm show that the proportion of species endemic to an island is positively related to its species richness and, assuming that endemism indexes speciation rate, they infer that greater species diversity accelerates diversification. Here we demonstrate that the same correlation between species richness and percentage endemism can arise even if within-island speciation is negligible, particularly when both endemism and species richness depend on attributes of islands (such as area) that influence the average age of resident populations. Island biogeography theory indicates that, where the average time to extinction is relatively long, diversity increases through colonization, irrespective of whether new species are formed; at the same time, islands on which populations persist for longer accumulate more endemic species as local populations differentiate and populations on neighbouring islands become extinct. We therefore suggest that species richness and endemism are correlated fortuitously owing to their mutual dependence on the life spans of populations on islands, which is unrelated to speciation itself.