Robert W. Elwood

Assessment of fighting ability in animal contests

Selection should favour accurate information gathering regarding the likely costs and benefits of continued conflict. Here we consider how variation in the abilities of contestants to assess resource-holding potential (RHP) influences fights. This has been examined in various game theory models. However, discriminating between assessment strategies has proven difficult and has resulted in confusion. To add clarity, we group existing models into three main types that differ in the information about RHP that contestants are presumed to gather: (1) pure self-assessment, (2) cumulative assessment and (3) mutual assessment. Within this framework we outline methods advocated to discriminate successfully between the three main assessment models. We discuss support for each model, before highlighting a number of conflicting and inconclusive studies, leading us to consider alternative approaches to investigate assessment. Furthermore, we examine support for newly emerging concepts such as ‘varying degrees of assessment’, ‘switching assessment’ strategies and the possibility of contestants adopting different assessment strategies within a fight involving distinctive roles. We suggest future studies will benefit by judicious use of a battery of techniques to determine how animals settle contests. Finally, we highlight difficulties with current game theory models, and raise concerns regarding the use of certain behavioural criteria to accept or reject a model, particularly since this may conflict with evidence for a given assessment strategy. Furthermore, the failure of existing models to account for newly emerging concepts points to limitations of their use and leads us to challenge game theoreticians to develop upon them.

THE TROPHIC ECOLOGY OF FRESHWATER GAMMARUS SPP. (CRUSTACEA:AMPHIPODA): PROBLEMS AND PERSPECTIVES CONCERNING THE FUNCTIONAL FEEDING GROUP CONCEPT

Gammarus spp. are widespread throughout a diverse range of freshwater habitats and can be the dominant part of many benthic macroinvertebrate assemblages, in terms of both numbers and/or biomass. Although the vast majority of studies have emphasized the herbivorous nature of Gammarus spp. and their ‘shredder’ functional feeding group (FFG) classification, we show that a far wider food base is exploited than has been previously acknowledged. This ‘plasticity’ as herbivore/predator is linked to the success of Gammarus spp. in persisting in and colonizing/invading disturbance‐prone ecosystems. Intraguild predation and cannibalism are more common than previously realized. This behaviour appears to be a causal mechanism in many amphipod species replacements. Additionally, Gammarus spp. are major predators of other members of the macroinvertebrate community. Furthermore, while many studies have emphasized fish predation on Gammarus spp., we illustrate how this fish: amphipod, predator: prey interaction may be a two‐way process, with Gammarus spp. themselves preying upon juvenile and wounded/trapped fish. We urge that a new realism be adopted towards the trophic ecology of Gammarus spp. and their role as predators and prey and that previously established FFG assumptions of both the food and the feeder be questioned critically.

The mismeasure of animal contests

Abstract Contests between rivals placing similar value on the resource at stake are commonly won by the rival having greater ‘resource holding potential’ (RHP). Mutual assessment of RHP difference between rivals is usually expected as an economical means of resolution; weaker rivals can retreat when they detect their relative inferiority, thereby avoiding costly, futile persistence. Models of contest resolution that entail retreat decisions based on estimates of RHP difference predict that contest duration diminishes as RHP difference between rivals increases because the asymmetry is more readily detected. This prediction appears to have been fulfilled in contests of diverse taxa, generating widespread support for assessment of RHP differences in contests. But few studies have considered alternatives in which each rival simply persists in accord with its own RHP (‘own RHP-dependent persistence’). In contests decided by own RHP-dependent persistence, in which costs accrue only through each rivals own actions, weaker rivals retreat first because they are inherently less persistent, and contest duration depends primarily on the weaker (losing) rivals RHP rather than RHP difference between the rivals. We show here that the analyses most commonly used to detect effects of RHP difference cannot discriminate between these alternatives. Because RHP difference between rivals tends to be correlated with RHP of the weaker rival in a pair, a negative relation between RHP difference and contest duration may be generated even when decisions of retreat are not based on estimated RHP difference. Many studies purporting to show a negative relation between RHP difference and contest duration may actually reflect an incidental association between weaker rival RHP and RHP difference. We suggest statistical and experimental approaches that may help to discriminate between effects of weaker rival RHP and true effects of RHP difference. We also discuss whether ‘true’ negative effects of RHP difference on contest duration always reflect retreat decisions based on estimated RHP differences. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.

Information gathering and decision making about resource value in animal contests

Contestants are predicted to adjust the cost of a fight in line with the perceived value of the resource and this provides a way of determining whether the resource has been assessed. An assessment of resource value is predicted to alter an animals motivational state and we note different methods of measuring that state. We provide a categorical framework in which the degree of resource assessment may be evaluated and also note limitations of various approaches. We place studies in six categories: (1) cases of no assessment, (2) cases of internal state such as hunger influencing apparent value, (3) cases of the contestants differing in assessment ability, (4) cases of mutual and equal assessment of value, (5) cases where opponents differ in resource value and (6) cases of particularly complex assessment abilities that involve a comparison of the value of two resources. We examine the extent to which these studies support game theory predictions and suggest future areas of research.

Behavioural trait assessment as a release criterion: boldness predicts early death in a reintroduction programme of captive-bred swift fox ( Vulpes velox )

Reintroduction of captive-bred animals is a key approach in conservation attempts for many endangered species, however, post-release survival is often low. Rearing conditions may be unlike those encountered upon release and the animals may not have had experiences necessary for survival in the wild. Animals may also habituate in captivity to stimuli that may pose a danger after release and/or there may be selection for behavioural traits, in particular reduced fearfulness, that may not be suited for the wild. Here, variation in boldness was assessed in captive-bred swift fox (Vulpes velox) and tested for influence on survival after release. Radio-tracked individuals that died in the 6 months following release were those judged previously as bold. In the presence of novel stimuli in captivity, they had left their dens more quickly, approached more closely to the stimuli and shown more activities indicating low fear than did those that survived. These individuals were less suited for release. Future selection of release-candidates on the basis of behavioural variation should enhance the success of reintroduction programmes.

Shell Wars: Assessment Strategies and the Timing of Decisions in Hermit Crab Shell Fights

The shell fighting behaviour of the hermit crab Pagurus bernhardus was observed under experimental conditions. The relative sizes of the crabs and shell quality of the larger crab influenced the probability of occurrence of a shell fight. These two factors along with the quality of the smaller crabs shell influenced the probability of an escalated fight occurring. During a shell fight, the attacker was able to assimilate information concerning the defenders shell and to compare it with the shell in possession. On the basis of this comparison the attacker decided whether or not to evict the defender and effect a shell change. The time that the attacker took in assessment of the opponents shell was influenced by a) the quality of the attackers own shell and b) the ease of discrimination between the qualities of the two shells. Thus decisions were easier in some instances than in others. The defending crab did not have access to information about the attackers shell and therefore could only estimate the quality of its own shell. This asymmetry of information experienced by the crabs ensured that the attacker decided the outcome of a shell fight.

The influence of litter size and parental behaviour on the development of Mongolian gerbil pups.

Abstract Undisturbed pairs of Mongolian gerbils were observed rearing litters of one, three or five pups. Pup behaviour and physical development were also monitored. Activities of both parents are influenced by litter size; some activities, e.g. sniff pups, being performed the most with large litters but others, e.g. nest-build, being performed the most with small litters. Pups from litters of three show the fastest behavioural and physical development. The presence of the male influences female activities and offspring development. Females nest-build less and sniff the pups less when the male is present. Pups reared with their father, however, open their eyes earlier and are behaviourally more advanced than are pups reared by the mother alone.

Changes in the responses of male and female gerbils (Meriones unguiculatus) towards test pups during the pregnancy of the female

Abstract Mongolian gerbils, maintained in monogamous pairs, were each offered a single test pup in order to assess changes in the paternal and the maternal responses during the females pregnancy. Naive males, housed with non-pregnant females, treat the pup as food, but they respond in a paternal manner when their mate becomes pregnant. Females eat pups except during very late pregnancy, when they show maternal responses. Experienced males never eat pups regardless of the reproductive state of the female. However, experienced females respond in the same manner as do naive females. Thus both parents are brought into a ‘maternal’ state prior to the birth of their offspring.

Replacement of the indigenous amphipod Gammarus duebeni celticus by the introduced G. pulex: differential cannibalism and mutual predation.

1. Following introductions in the 1950s, Gammarus pulex has replaced the native Irish freshwater amphipod G. duebeni celticus in several river systems. We show that G. pulex has eliminated and replaced G. d. celticus in lower sections of the River Lagan system, N. Ireland, although G. d. celticus maintains pure populations upstream from G. pulex. More rarely, the two species may be found together. 2. We investigate the potential roles of cannibalism and mutual predation in explaining these replacement patterns. G. d. celticus was significantly more cannibalistic on moulted conspecifics than G. pulex

The dynamics of predation on gammarus spp. (Crustacea: Amphipoda)

Gammarus spp. (Crustacea: Amphipoda) are widespread throughout a diverse range of marine, freshwater and estuarine/brackish habitats, often dominating benthic macroinvertebrate communities in terms of both numbers and/or biomass. Gammarus spp. are the dominant macroinvertebrate prey items of many fish, whether as a seasonal food source or a year‐round staple. Selective predation by fish on Gammarus spp. is often linked to parasitism and the body size of the prey. Gammarus spp. populations are under increasing threat from both pollution and replacement/displacement by introduced species. Loss of populations and species invasions/replacements could have significant impacts on native predator species if the predator(s) cannot successfully adapt their feeding patterns to cope with non‐indigenous Gammarus prey species. Despite this, many fish predation studies do not identify Gammarus prey to species level. This lack of precision could be important, as Gammarus spp. exhibit wide variations in physiochemical tolerances, habitat requirements, abilities to invade and susceptibility to replacement. Although rarely acknowledged, the impacts of nonpiscean predators (particularly macroinvertebrates) on Gammarus prey species may frequently be stronger than those exerted by fish. A major aim of this review is to ascertain the current importance of Gammarus as a prey species, such that the implications of changes in Gammarus spp. populations can be more accurately assessed by interested groups such as ecologists and fisheries managers. We also review the dynamics of Gammarus spp. as prey to a diverse array of mammals, birds, amphibians, insects, flatworms, other crustaceans such as crabs and crayfish and, perhaps most importantly, other Gammarus spp.