Predation risk and resource abundance mediate foraging behaviour and intraspecific resource partitioning among consumers in dominance hierarchies

Dominance hierarchies and the resulting unequal resource partitioning among individuals are key mechanisms of population regulation. The strength of dominance hierarchies can be influenced by size‐dependent tradeoffs between foraging and predator avoidance whereby competitively inferior subdominants can access a larger proportion of limiting resources by accepting higher predation risk. Foraging‐predation risk tradeoffs also depend on resource abundance. Yet, few studies have manipulated predation risk and resource abundance simultaneously; consequently, their joint effect on resource partitioning within dominance hierarchies are not well understood. We addressed this gap by measuring behavioural responses of masu salmon Oncorhynchus masou ishikawae to experimental manipulations of predation risk and resource abundance in a natural temperate forest stream. Responses to predation risk depended on body size and social status such that larger fish (often social dominants) exhibited more risk‐averse behaviour (e.g. lower foraging and appearance rates) than smaller subdominants after exposure to a simulated predator. The magnitude of this effect was lower when resources were elevated, indicating that dominant fish accepted a higher predation risk to forage on abundant resources. However, the influence of resource abundance did not extend to the population level, where predation risk altered the distribution of foraging attempts (a proxy for energy intake) from being skewed towards large individuals to being skewed towards small individuals after predator exposure. Our results imply that size‐dependent foraging–predation risk tradeoffs can weaken the strength of dominance hierarchies by allowing competitively inferior subdominants to access resources that would otherwise be monopolized.     

Diseased Social Predators

Social predators benefit from cooperation in the form of increased hunting success, but may be at higher risk of disease infection due to living in groups. Here, we use mathematical modeling to investigate the impact of disease transmission on the population dynamics benefits provided by group hunting. We consider a predator–prey model with foraging facilitation that can induce strong Allee effects in the predators. We extend this model by an infectious disease spreading horizontally and vertically in the predator population. The model is a system of three nonlinear differential equations. We analyze the equilibrium points and their stability as well as one- and two-parameter bifurcations. Our results show that weakly cooperating predators go unconditionally extinct for highly transmissible diseases. By contrast, if cooperation is strong enough, the social behavior mediates conditional predator persistence. The system is bistable, such that small predator populations are driven extinct by the disease or a lack of prey, and large predator populations survive because of their cooperation even though they would be doomed to extinction in the absence of group hunting. We identify a critical cooperation level that is needed to avoid the possibility of unconditional predator extinction. We also investigate how transmissibility and cooperation affect the stability of predator–prey dynamics. The introduction of parasites may be fatal for small populations of social predators that decline for other reasons. For invasive predators that cooperate strongly, biocontrol by releasing parasites alone may not be sufficient.     

Two blue tit Parus caeruleus populations from Corsica differ in social dominance

Although the causes and consequences of social dominance have been examined extensively, avian studies have rarely focused on between-population differences in social dominance. On the island of Corsica, two resident blue tit Parus caeruleus populations 25 km apart differ significantly in body size measures, timing and effort of reproduction, and song structure, and some of these population differences have a genetic basis. Because earlier avian studies have shown that social dominance is influenced by body size or mass, we predicted that individuals from these two blue tit populations would also differ in their ability to dominate other individuals. Consistent with this prediction, we found that male blue tits of these two populations differ in social dominance, and that heavier or larger individuals dominate lighter or smaller ones in aviary experiments. We propose that social dominance may serve to maintain phenotypic population differentiation at a micro-geographic scale by acting as a barrier to dispersal.     

On dominance relations and the structure of animal societies: II. Some effects of possible social factors

In a previous paper (Landau, 1951) it was shown that a society with a dominance relation would rarely tend to be close to the hierarchy in structure if dominance is determined solely by the inherent characteristics of the members. Here we consider the effects of other factors, due to social rank or to the outcome of previous encounters which affected dominance. The following results are obtained. A uniform bias against reversal of dominance will have no effect on the stationary distribution of the structure of the society. If the probability of dominance is a linear function of the previously established score (number of members dominated), there will be a small tendency for the society to move toward the hierarchy; but this is negligible for large societies. If a member never challenges another whose score exceeds his own by two or more, or if he can never dominate if he should challenge, then the hierarchy is the only stable structure. From the last result it is concluded that social factors which restrict challenges or the probability of dominance could easily account for societies close to the hierarchy, such as are observed in flocks of domestic hens. The effectiveness of social bias in establishing hierarchies is much greater in small societies than in large ones.     

Self-organizing dominance hierarchies in a wild primate population

Linear dominance hierarchies, which are common in social animals, can profoundly influence access to limited resources, reproductive opportunities and health. In spite of their importance, the mechanisms that govern the dynamics of such hierarchies remain unclear. Two hypotheses explain how linear hierarchies might emerge and change over time. The ‘prior attributes hypothesis’ posits that individual differences in fighting ability directly determine dominance ranks. By contrast, the ‘social dynamics hypothesis’ posits that dominance ranks emerge from social self-organization dynamics such as winner and loser effects. While the prior attributes hypothesis is well supported in the literature, current support for the social dynamics hypothesis is limited to experimental studies that artificially eliminate or minimize individual differences in fighting abilities. Here, we present the first evidence supporting the social dynamics hypothesis in a wild population. Specifically, we test for winner and loser effects on male hierarchy dynamics in wild baboons, using a novel statistical approach based on the Elo rating method for cardinal rank assignment, which enables the detection of winner and loser effects in uncontrolled group settings. Our results demonstrate (i) the presence of winner and loser effects, and (ii) that individual susceptibility to such effects may have a genetic basis. Taken together, our results show that both social self-organization dynamics and prior attributes can combine to influence hierarchy dynamics even when agonistic interactions are strongly influenced by differences in individual attributes. We hypothesize that, despite variation in individual attributes, winner and loser effects exist (i) because these effects could be particularly beneficial when fighting abilities in other group members change over time, and (ii) because the coevolution of prior attributes and winner and loser effects maintains a balance of both effects.     

Feedback between size balance and consumption strongly affects the consequences of hatching phenology in size‐dependent predator–prey interactions

In many size‐dependent predator–prey systems, hatching phenology strongly affects predator–prey interaction outcomes. Early‐hatched predators can easily consume prey when they first interact because they encounter smaller prey. However, this process by itself may be insufficient to explain all predator–prey interaction outcomes over the whole interaction period because the predator–prey size balance changes dynamically throughout their ontogeny. We hypothesized that hatching phenology influences predator–prey interactions via a feedback mechanism between the predator–prey size balance and prey consumption by predators. We experimentally tested this hypothesis in an amphibian predator–prey model system. Frog tadpoles Rana pirica were exposed to a predatory salamander larva Hynobius retardatus that had hatched 5, 12, 19 or 26 days after the frog tadpoles hatched. We investigated how the salamander hatch timing affected the dynamics of prey mortality, size changes of both predator and prey, and their subsequent life history (larval period and size at metamorphosis). The predator–prey size balance favoured earlier hatched salamanders, which just after hatching could successfully consume more frog tadpoles than later hatched salamanders. The early‐hatched salamanders grew rapidly and their accelerated growth enabled them to maintain the predator‐superior size balance; thus, they continued to exert strong predation pressure on the frog tadpoles in the subsequent period. Furthermore, frog tadpoles exposed to the early‐hatched salamanders were larger at metamorphosis and had a longer larval period than other frog tadpoles. These results suggest that feedback between the predator‐superior size balance and prey consumption is a critical mechanism that strongly affects the impacts of early hatching of predators in the short‐term population dynamics and life history of the prey. Because consumption of large nutrient‐rich prey items supports the growth of predators, a similar feedback mechanism may be common and have strong impacts on phenological shifts in size‐dependent trophic relationships.     

Metamorphosing reef fishes avoid predator scent when choosing a home

Most organisms possess anti-predator adaptations to reduce their risk of being consumed, but little is known of the adaptations prey employ during vulnerable life-history transitions when predation pressures can be extreme. We demonstrate the use of a transition-specific anti-predator adaptation by coral reef fishes as they metamorphose from pelagic larvae to benthic juveniles, when over half are consumed within 48 h. Our field experiment shows that naturally settling damselfish use olfactory, and most likely innate, predator recognition to avoid settling to habitat patches manipulated to emit predator odour. Settlement to patches emitting predator odour was on average 24-43% less than to control patches. Evidence strongly suggests that this avoidance of sedentary and patchily distributed predators by nocturnal settlers will gain them a survival advantage, but also lead to non-lethal predator effects: the costs of exhibiting anti-predator adaptations. Transition-specific anti-predator adaptations, such as demonstrated here, may be widespread among organisms with complex life cycles and play an important role in prey population dynamics.     

Female philopatry and dominance patterns in wild geladas

Cercopithecines have a highly conserved social structure with strong female bonds and stable, maternally inherited linear dominance hierarchies. This system has been ascribed to the pervasiveness of female philopatry within the typical multi-male, multi-female social groups. We examined the relationship between female philopatry, dominance hierarchies, and reproduction in geladas (Theropithecus gelada), a species with an unusual multi-leveled society. During a 4-year field study on a wild population in the Simien Mountains National Park, Ethiopia, we observed 14 units across two bands of geladas that underwent a number of events, such as male takeovers and female deaths, which could potentially disrupt female relationships and unit structure. First, we corroborate earlier reports that gelada females are natally philopatric: we observed no interunit migrations, and the female mortality rate was comparable to that of philopatric baboons (suggesting all female disappearances were indeed deaths). Second, contrary to previous reports, data from this long-term study show that geladas exhibit the linear and stable dominance hierarchies typical of other Cercopithecines. Moreover, female ranks appear to be maternally inherited. Third, we found no evidence that alpha females aggressively target the lowest ranking individuals, nor did rank confer clear reproductive advantages to dominant females within our 4-year observation period. As such, geladas fit the allostatic load model [Goymann & Wingfield, Animal Behaviour 67:591-602, 2004]. Our study confirms the importance of female philopatry in the kin-based Cercopithecine dominance system.     

Spatial clumping of food and social dominance affect interference competition among ruddy turnstones

In studying the success of foraging animals, studies of interferencecompetition have put emphasis on effects of competitor density,whereas studies of resource defense have focused on the effectsof the spatial distribution of food within patches. Very fewstudies have looked at both factors simultaneously, that is,determined whether the effects of competitor density on foragingsuccess depend on the spatial distribution of food. We studiedthe behavior and the foraging success of ruddy turnstones (Arenariainterpres) using an experiment in which we varied both the presenceof a competitor and the food distribution. Because turnstonesmay differ strongly in their relative dominance status, we alsoexperimentally varied the foragers' relative dominance status.We found that the presence of a competitor only reduced theforaging success of subordinate birds foraging at the clumpedfood distribution. At this condition, dominant and subordinatebirds differed markedly in their foraging success. Contraryto our expectations, we did not observe more agonistic behaviorat the clumped food distribution. This indicates that the amountof agonistic behavior observed may be a bad indicator of interferenceeffects. These findings have specific implications for modelsof interference competition. Most notably they show that theeffects of competitor density on agonistic behavior and foragingsuccess may well depend on the spatial distribution of foodand the foragers' relative dominance status. Additionally, ourresults suggest that social dominance will not be fully understoodwithout considering long-term processes such as the formationand maintenance of social dominance hierarchies.     

Co-evolution and ecosystem based problem solving

Emergent cooperative relations in ecosystems are ill understood, but have the potential to strongly improve evolutionary computing. On the other hand, eco-evolutionary computation has the potential to provide new insights in the structuring and functioning of ecosystems. Here we study ecosystem based problem solving in a co-evolutionary framework of predators (solvers) and prey (problems), extended with a population of scavengers, which can eat the remains of prey (that is, cooperate with the predators in solving the problems). We show that such an artificial ecosystem of predators, prey and scavengers, with a selection and fitness regime favoring specialization, self-organizes in space and time such that (1) problems are automatically decomposed in easier to solve parts, (2) the predator, prey and scavenger populations differentiate in sub-populations according to this decomposition, and (3) predators and scavengers automatically co-localize in space such that the problems are indeed solved by predator–scavenger combinations which together correctly approximate the target function. That is, the use of a spatial co-evolutionary ecosystem as information processing unit for evolutionary computation gives rise to an emergent structure of niches, each consisting of complementary partial solutions. As a result, ecosystem based solutions are preferred over individual-based solutions in solving the studied function approximation task.     

Innate threat-sensitive foraging: black-tailed deer remain more fearful of wolf than of the less dangerous black bear even after 100 years of wolf absence

Anti-predator behaviors often entail foraging costs, and thus prey response to predator cues should be adjusted to the level of risk (threat-sensitive foraging). Simultaneously dangerous predators (with high hunting success) should engender the evolution of innate predator recognition and appropriate anti-predator behaviors that are effective even upon the first encounter with the predator. The above leads to the prediction that prey might respond more strongly to cues of dangerous predators that are absent, than to cues of less dangerous predators that are actually present. In an applied context this would predict an immediate and stronger response of ungulates to the return of top predators such as wolves (Canis lupus) in many parts of Europe and North America than to current, less threatening, mesopredators. We investigated the existence of innate threat-sensitive foraging in black-tailed deer. We took advantage of a quasi-experimental situation where deer had not experienced wolf predation for ca. 100 years, and were only potentially exposed to black bears (Ursus americanus). We tested the response of deer to the urine of wolf (dangerous) and black bear (less dangerous). Our results support the hypothesis of innate threat-sensitive foraging with clear increased passive avoidance and olfactory investigation of cues from wolf, and surprisingly none to black bear. Prey which have previously evolved under high risk of predation by wolves may react strongly to the return of wolf cues in their environments thanks to innate responses retained during the period of predator absence, and this could be the source of far stronger non-consumptive effects of the predator guild than currently observed.     

Hierarchical structure and the influence of individual attributes in the captive squirrel monkey (<Emphasis Type="Italic">Saimiri collinsi</Emphasis>)

The dominance structure of primate social groups varies widely. In addition to the groups’ composition, intrinsic attributes such as sex, body size and life experience are important factors that can affect hierarchical dominance relations. All primates are social animals, and the social environment has a direct influence on the physiological conditions of vital systems such as immunological, reproductive and cardiovascular systems. In this study, we analyze the hierarchical structure of Saimiri collinsi in captivity, including the hierarchical structure type, the influence of individual intrinsic characteristics (sex, age, weight and origin—born in captivity or in the wild) based on the prior-attributes model, the relation between agonistic behavior frequency and hierarchical position, and hierarchy steepness, which represents the dominance gradient. We found that the group order was characterized by a partial hierarchy: a dominance position could be occupied by more than one individual simultaneously, including individuals of both sexes. Intrinsic characteristics had no influence on hierarchical structure, with the exception of the male in the highest hierarchical position, which had a markedly larger body than all other group members. Thus, the prior-attributes model did not apply to hierarchical formation of S. collinsi in captivity. Only the frequency of agonistic behavior of males correlated with their hierarchical position, and they differed from all other group members in their more aggressive behavior. The steepness between adjacent positions along the dominance gradient was significant only between the dominant male and the next individual in the group, with a smooth gradient between the other positions in the rank. As the access to resources is directly related to hierarchical dominance, a smooth dominance gradient is to be expected in species that form very large groups, such as wild Saimiri populations.     

Ecological determinants of herd size in the Thornicroft’s giraffe of Zambia

Ecological factors have a pervasive impact on animal population sizes and the structure of their social systems. In a number of ungulate species, predator pressure exerts a major influence on group size. Given that giraffe (Giraffa camelopardalis) live in an extremely flexible social system, and that breeding is nonseasonal, they are an ideal species for examining how ecological variables contribute to fluctuations in herd size. We present an analysis of 34 years of data on a population of Thornicroft’s giraffe (G. c. thornicrofti Lydekker 1911) that reveal how herd size changes with season and habitat. Sex differences in herd size were apparent, with bulls often travelling as singletons, whereas cows were generally observed with conspecifics. Herds were larger during the wet than dry season, but herd size changed in a parallel fashion across habitats. Giraffe herds were smaller in woodland and thicket areas than in open habitats, regardless of season. We suggest that the regular fluctuations in herd size among giraffe indicate a fission/fusion social system embedded within a larger social community. We conclude that changes in herd size among giraffe reflect a dynamic process regulated by individuals adjusting the number of associates based upon an interaction of foraging, reproductive, social and antipredator strategies.     

Herbivore and predator diversity interactively affect ecosystem properties in an experimental marine community

Interacting changes in predator and prey diversity likely influence ecosystem properties but have rarely been experimentally tested. We manipulated the species richness of herbivores and predators in an experimental benthic marine community and measured their effects on predator, herbivore and primary producer performance. Predator composition and richness strongly affected several community and population responses, mostly via sampling effects. However, some predators survived better in polycultures than in monocultures, suggesting complementarity due to stronger intra- than interspecific interactions. Predator effects also differed between additive and substitutive designs, emphasizing that the relationship between diversity and abundance in an assemblage can strongly influence whether and how diversity effects are realized. Changing herbivore richness and predator richness interacted to influence both total herbivore abundance and predatory crab growth, but these interactive diversity effects were weak. Overall, the presence and richness of predators dominated biotic effects on community and ecosystem properties.     

Revealing the role of predator interference in a predator–prey system with disease in prey population

Predation on a species subjected to an infectious disease can affect both the infection level and the population dynamics. There is an ongoing debate about the act of managing disease in natural populations through predation. Recent theoretical and empirical evidence shows that predation on infected populations can have both positive and negative influences on disease in prey populations. Here, we present a predator–prey system where the prey population is subjected to an infectious disease to explore the impact of predator on disease dynamics. Specifically, we investigate how the interference among predators affects the dynamics and structure of the predator–prey community. We perform a detailed numerical bifurcation analysis and find an unusually large variety of complex dynamics, such as, bistability, torus and chaos, in the presence of predators. We show that, depending on the strength of interference among predators, predators enhance or control disease outbreaks and population persistence. Moreover, the presence of multistable regimes makes the system very sensitive to perturbations and facilitates a number of regime shifts. Since, the habitat structure and the choice of predators deeply influence the interference among predators, thus before applying predators to control disease in prey populations or applying predator control strategy for wildlife management, it is essential to carefully investigate how these predators interact with each other in that specific habitat; otherwise it may lead to ecological disaster.     

Testosterone and Linear Social Dominance Status in Captive Male Dabbling Ducks in Winter

Dominance hierarchies play an important role in avoidance and/or solving conflicts in gregarious species. In dabbling ducks (Anas species), dominance allows for feeding‐site monopolization in winter quarters where resources are generally limited. In addition, male social rank should theoretically favour access to mates. Dominance rank can be associated with morphological traits, and is often correlated with aggressiveness, a behavioural trait generally related to high testosterone levels. In this study, we investigated the existence of a winter group structure based on dominance relationships and tested for a linear hierarchy, in three species of captive male dabbling ducks (mallard Anas platyrhynchos, pintail A. acuta and wigeon A. penelope). We then analysed the relationship between dominance ranks, morphological parameters and testosterone levels measured in early (Oct.) and mid‐winter (Dec./Jan.). We found that the three male groups of the three species exhibited a linear hierarchy. Testosterone levels differed during winter and between species. Morphologic measurements, body mass and body condition were not correlated with individual dominance ranks, whereas dominant males had higher testosterone levels than subordinates. The slopes of the relationships were similar between species and winter period, but the y‐intercepts differed between species and between early and mid‐winter phases. The linear hierarchy found in the three species indicates that dominance relationships strongly structure dabbling duck groups in winter. Lack of correlation between rank and morphological characters, but correlation of rank with testosterone levels suggests that social rank is more dependent on behavioural traits such as aggressive behaviour. The differences between species and winter periods are discussed in relation to migration and wintering phenology.     

Allee效应对具有生境恢复的集合种群的影响

Allee效应与种群的灭绝密切相关,其研究对生态保护和管理至关重要。Allee效应对物种续存是潜在的干扰因素,濒危物种更容易受其影响,可能会增加生存于生境破碎化斑块的濒危物种的死亡风险,因此研究Allee效应对种群的动态和续存的影响是必要的。从包含由生物有机体对环境的修复产生的Allee效应的集合种群模型出发,引入由其他机制形成的Allee效应,建立了常微分动力系统模型和基于网格模型的元胞自动机模型。通过理论分析和计算机模拟表明:(1)强Allee效应不利于具有生境恢复的集合种群的续存;(2)生境恢复有利于种群续存;(3)局部扩散影响了集合种群的空间结构、动态行为和稳定性,生境斑块之间的局部作用将会减缓或消除集合种群的Allee效应,有利于集合种群的续存。     

Relation between complexity and stability in food webs with adaptive behavior

We investigate the influence of functional responses (Lotka-Volterra or Holling type), initial topological web structure (randomly connected or niche model), adaptive behavior (adaptive foraging and predator avoidance) and the type of constraints on the adaptive behavior (linear or nonlinear) on the stability and structure of food webs. Two kinds of stability are considered: one is the network robustness (i.e., the proportion of species surviving after population dynamics) and the other is the species deletion stability. When evaluating the network structure, we consider link density as well as the trophic level structure. We show that the types of functional responses and initial web structure do not have a large effect on the stability of food webs, but foraging behavior has a large stabilizing effect. It leads to a positive complexity-stability relationship whenever higher "complexity" implies more potential prey per species. The other type of adaptive behavior, predator avoidance behavior, makes food webs only slightly more stable. The observed link density after population dynamics depends strongly on the presence or absence of adaptive foraging, and on the type of constraints used. We also show that the trophic level structure is preserved under population dynamics with adaptive foraging.     

On dominance relations and the structure of animal societies: I. Effect of inherent characteristics

Societies are considered in which a non-transitive dominance relation exists between every pair of members, such as the peck-right in a flock of hens. A one-dimensional measure of the structure of such a society,h, is defined, withh=0 for equality andh=1 for the hierarchy. It is assumed that each member of the society is characterized by an ability vector whose components depend on individual characteristics such as size, concentration of sex hormone, etc., but not on social factors such as social rank. The distribution of abilities among members of the society is assumed to be given by a distribution function which is the same for all members, and the probability that one member dominates another is given by a function of the ability vectors of the two. On these assumptions formulas for the expected (mean) value and variance ofh are determined in terms of the distribution and dominance probability functions. Some special cases are calculated, especially that for normany distributed abilities and dominance probability given by the normal probability integral. Several conclusions are derived. If all members are of equal ability, so that dominance probability is 1/2, then any sizable society is much more likely to be near the equality than the hierarchy; and, as the size of the society increases, the probability that it will be near the hierarchy becomes vanishingly small. If the dominance probability is a weighted sum of several independent components, which make up the ability vector, then the society is less likely to be close to the hierarchy as the number of these components increases. The hierarchy is the prevalent structure only if unreasonably small differences in ability are decisive for dominance. From this it appears that the social factors, or psychological factors such as the previous history of dominance, which are not included in the present treatment, may be of great importance in explaining the observed prevalence of structures very close to the hierarchy in flocks of domestic hens.     

Context-dependent effects of predator removal from experimental microcosm communities

The loss of a predator from an ecological community can cause large changes in community structure and ecosystem processes, or have very little consequence for the remaining species and ecosystem. Understanding when and why the loss of a predator causes large changes in community structure and ecosystem processes is critical for understanding the functional consequences of biodiversity loss. We used experimental microbial communities to investigate how the removal of a large generalist predator affected the extinction frequency, population abundance and total biomass of its prey. We removed this predator in the presence or absence of an alternative, more specialist, predator in order to determine whether the specialist predator affected the outcome of the initial species removal. Removal of the large generalist predator altered some species' populations but many were unaffected and no secondary extinctions were observed. The specialist predator, though rare, altered the response of the prey community to the removal of the large generalist predator. In the absence of the specialist predator, the effects of the removal were only measurable at the level of individual species. However, when the specialist predator was present, the removal of the large generalist predator affected the total biomass of prey species. The results demonstrate that the effect of species loss from high trophic levels may be very context-dependent, as rare species can have disproportionately large effects in food webs.