Costs increase as ritualized fighting progresses within and between phases in the sierra dome spider, Neriene litigiosa

Magnitudes and patterns of energy expenditure in animal contests are seldom measured, but can be critical for predicting contest dynamics and understanding the evolution of ritualized fighting behaviour. In the sierra dome spider, males compete for sexual access to females and their webs. They show three distinct phases of fighting behaviour, escalating from ritualized noncontact display (phase 1) to cooperative wrestling (phase 2), and finally to unritualized, potentially fatal fighting (phase 3). Using CO2 respirometry, we estimated energetic costs of male-male combat in terms of mean and maximum metabolic rates and the rate of increase in energy expenditure. We also investigated the energetic consequences of age and body mass, and compared fighting metabolism to metabolism during courtship. All three phases involved mean energy expenditures well above resting metabolic rate (3.5 X, 7.4 X and 11.5 X). Both mean and maximum energy expenditure became substantially greater as fights escalated through successive phases. The rates of increase in energy use during phases 2 and 3 were much higher than in phase 1. In addition, age and body mass affected contest energetics. These results are consistent with a basic prediction of evolutionarily stable strategy contest models, that sequences of agonistic behaviours should be organized into phases of escalating energetic costs. Finally, higher energetic costs of escalated fighting compared to courtship provide a rationale for first-male sperm precedence in this spider species. (C) 2004 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Human behaviour modelled as a statistico-deterministic turing machine

'Free will' and its corollary, the concept of individual responsibility are keystones of the justice system. This paper shows that if we accept a physics that disallows time reversal, the concept of 'free will' is undermined by an integrated understanding of the influence of genetics and environment on human behavioural responses. Analysis is undertaken by modelling life as a novel statistico-deterministic version of a Turing machine, i.e. as a series of transitions between states at successive instants of time. Using this model it is proven by induction that the entire course of life is independent of the action of free will. Although determined by prior state, the probability of transitions between states in response to a standard environmental stimulus is not equal to 1 and the transitions may differ quantitatively at the molecular level and qualitatively at the level of the whole organism. Transitions between states correspond to behaviours. It is shown that the behaviour of identical twins (or clones), although determined, would be incompletely predictable and non-identical, creating an illusion of the operation of 'free will'. 'Free will' is a convenient construct for current judicial systems and social control because it allows rationalization of punishment for those whose behaviour falls outside socially defined norms. Indeed, it is conceivable that maintenance of ideas of free will has co-evolved with community morality to reinforce its operation. If the concept is free will is to be maintained it would require revision of our current physical theories.

How primatology can inform us about the evolution of the humand mind

Humans are primates. We have evolved from common ancestors and the evolution of the human body is becoming increasingly clear as the archeological record expands. But for most people the gap between humans and animals lies in the mind, not in the body. And minds do not fossilise. To reconstruct the evolution of mind, scholars have thus increasingly looked to our closest relatives for clues. Here I discuss four ways in which the study of primates may inform such reconstruction: fact-finding, phylogenetic reconstruction, analogy, and regression models. Knowledge about primates can help us bridge the gap. Extinction of our closest relatives, on the other hand, would not only deplete that source of information but also increase the apparent differences between animal and human minds. It is likely that we have a long history of displacing closely related species, including the other hominids, leading us to appear ever more unique.

Imitation recognition in a captive chimpanzee (pan troglodytes)

This study investigated the ability of a captive chimpanzee (Pan troglodytes) to recognise when he is being imitated. In the experimental condition of test 1a, an experimenter imitated the postures and behaviours of the chimpanzee as they were being displayed. In three control conditions the same experimenter exhibited (1) actions that were contingent on, but different from, the actions of the chimpanzee, (2) actions that were not contingent on, and different from, the actions of the chimpanzee, or (3) no action at all. The chimpanzee showed more "testing" sequences (i.e., systematically varying his actions while oriented to the imitating experimenter) and more repetitive behaviour when lie was being imitated, than when he was not. This finding was replicated 4 months later in test 1b. When the experimenter repeated the same actions she displayed in the experimental condition of test 1a back to the chimpanzee in test 2, these actions now did not elicit those same testing sequences or repetitive behaviours. However, a live imitation condition did. Together these results provide the first evidence of imitation recognition in a nonhuman animal.

Communication in coral reef fish: the role of ultraviolet colour patterns in damselfish territorial behaviour

Many coral reef fish possess ultraviolet (UV) colour patterns. The behavioural significance of these patterns is poorly understood and experiments on this issue have not been reported for free-living reef fish in their natural environment. The damselfish Pomacentrus amboinensis has UV facial patterns, and spectroradiometric ocular media measurements show that it has the potential for UV vision. To test the potential behavioural significance of the UV patterns, I studied the response of males, in natural territories on the reef and in aquaria, to two conspecific intruders, one presented in a UV-transmitting (UV+) container and the other in a UV-absorbing (UV-) one. Territory owners attacked intruders viewed through UV+ filters significantly more often and for longer than intruders viewed through the UV- filter. In general, the results of the field experiment confirmed those of the laboratory experiment. The results support the hypothesis that P. amboinensis males are sensitive to UV light and that reflectance patterns, which appear in high contrast only in UV, modulate the level of aggressive behaviour. A recent survey showed that many predatory fish may not have UV vision and the use of UV colours in select species of reef fish may therefore serve as a 'private communication channel'. (C) 2004 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Contrasting mutual sexual selection on homologous signal traits in Drosophila serrata

The nature of male mating preferences, and how they differ from female mating preferences in species with conventional sex roles, has received little attention in sexual selection studies. We estimated the form and strength of sexual selection as a consequence of male and female mating preferences in a laboratory-based population of Drosophila serrata. The differences between sexual selection on male and female signal traits (cuticular hydrocarbons [CHCs]) were evaluated within a formal framework of linear and nonlinear selection gradients. Females tended to exert linear sexual selection on male CHCs, whereas males preferred intermediate female CHC phenotypes leading to convex (stabilizing) selection gradients. Possible mechanisms determining the nonlinear nature of sexual selection on female CHCs are proposed.

Foraging behaviour of Helicoverpa armigera first instar larvae on crop plants of different developmental stages

Understanding how insect pests forage on their food plants can help optimize management strategies. Helicoverpa armigera (Hubner) (Lep., Noctuidae) is a major polyphagous pest of agricultural crops worldwide. The immature stages feed and forage on crops at all stages of plant development, damaging fruiting and non-fruiting structures, yet very little is known about the influence of host type or stage on the location and behaviour of larvae. Through semi-continuous observation, we evaluated the foraging (movement and feeding) behaviours of H. armigera first instar larvae as well as the proportion of time spent at key locations on mungbean [Vigna radiata (L.) Wilczek] and pigeon pea [Cajanus cajan (L.) Millspaugh] of differing developmental stages: seedling- and mature (flowering/pod fill)-stage plants. Both host type and age affected the behaviour of larvae. Larvae spent more time in the upper parts of mature plants than on seedlings and tended to stay at the top of mature plants if they moved there. This difference was greater in pigeon pea than in mungbean. The proportion of time allocated to feeding on different parts of a plant differed with host and age. More feeding occurred in the top of mature pigeon pea plants but did not differ between mature and seedling mungbean plants. The duration of key behaviours did not differ between plant ages in either crop type and was similar between hosts although resting bouts were substantially longer on mungbeans. Thus a polyphagous species such as H. armigera does not forage in equivalent ways on different hosts in the first instar stage.

Geographic variation in vocalisations of Satin Bowerbirds, Ptilonorynchus violaceus, in south-eastern Queensland

Geographic variation in the advertisement call of the male Satin Bowerbird, Ptilonorhynchus violaceus, was investigated in three populations in south-eastern Queensland. The call was found to differ significantly among the three geographically distinct populations. A discriminant function analysis using five measurements of call frequency and duration provided 100% classification success of the 25 individuals. The observed geographic variation in this call may result from adaptation to the local acoustic environment in these populations, or from genetic or cultural divergence among populations. Further research involving the acoustic properties of the habitats, population genetics and a larger number of populations is required to fully understand this pattern of call variation.

The multiple signals assessed by female satin bowerbirds: could they be used to narrow down females' choices of mates?

Female choice based on multiple male traits has been documented in many species but the functions of such multiple traits are still under debate. The satin bowerbird has a polygynous mating system in which males attract females to bowers for mating; females choose mates based on multiple aspects of males and their bowers. In this paper, we demonstrate that females use some cues to decide which males to examine closely and other cues to decide which males to mate with. Female visitation rates to bowers were significantly related to male size and the males' 'solitary' display rates, and, to a lesser extent, to the numbers of bower decorations. After controlling for female visitation rates, it was found that a male's mating success was significantly related to his size and the rate at which he 'painted' his bower with saliva and chewed up plant material.

Influence of Observer Presence on Pacific Damsel Bug Behavior: Who is Watching Whom?

When direct observations are used to study animal behavior the presence of the observer may alter the animal and hence influence the data being collected, yet few studies have quantified this effect. We conducted direct observation studies in the glasshouse to assess the relative influence of observer presence on the behavior of Pacific damsel bugs, Nabis kinbergii, a potentially important predator of crop pests. Comparisons of predator activity, predator distribution, prey (Helicoverpa armigera) mortality and prey distribution between frequently observed and minimally observed treatments, during diurnal and nocturnal observation sessions showed that the frequency of observer presence had no apparent impact on Pacific damsel bug behavior. This is the first documented test of the impact of observer presence in an insect system. To place our results in context, we reviewed 15 papers on the influence of observer presence in a range of animals. We established that just over half of these papers found evidence for an effect. Nevertheless, direct observations should be useful in further studies of Pacific damsel bug behavior, and researchers using direct observations to study the behavior of other animals should be cognizant of observer effects during design and interpretation of their study.

Factors affecting the vigilance and flight behaviour of impalas

This study investigated the influences of various natural and anthropogenic factors on the vigilance and flight behaviour of impalas in the Save Valley Conservancy, Zimbabwe, using multivariate statistical techniques. The factor that most significantly affected the proportions of time that individuals spent being vigilant and their rates of vigilance was the position of a focal animal in the group; impalas on the periphery of a group were more vigilant than central impalas. Both measures of vigilance were also negatively related to group size. Males spent more total time being vigilant but females raised their heads more often. Impalas spent more time being vigilant in the late afternoon than in the early morning, when greater than ten metres from cover, and when predators had been nearby within the previous six hours. Impalas spent more time vigilant at the property where more impalas were hunted, possibly reflecting the differences in the intensity of hunting by humans on the two properties. Flight distances at the approach of humans were significantly greater at one property than the other, and were also greater for small groups. Further research into the effects of hunting by humans on animals' antipredator behaviours would provide valuable insights for wildlife managers.

Moreton Bay, Queensland, Australia: An example of the co-existence of significant marine mammal populations and large-scale coastal development

Recent analyses assert that large marine vertebrates such as marine mammals are now 'functionally or entirely extinct in most coastal ecosystems'. Moreton Bay is a large diverse marine ecosystem bordering the fastest growing area in Australia. The human population is over 1.6 million and increasing yearly by between 10% and 13% with resultant impacts upon the adjoining marine environment. Nonetheless, significant populations of three species of marine mammals are resident within Moreton Bay and a further 14 species are seasonal or occasional visitors. This paper reviews the current and historical distributions and abundance of these species in the context of the current management regime and suggests initiatives to increase the resilience of marine mammal populations to the changes wrought by the burgeoning human population in coastal environments. (C) 2004 Elsevier Ltd. All rights reserved.

Division of labour within cooperatively breeding groups

Within cooperative societies, group members share in caring for offspring. Although division of labour among group members has been relatively well studied in insects, less is known about vertebrates. Most studies of avian helping focus solely on the extent to which helpers provision the offspring, however, helpers can participate in everything from nest building to predator defence. Bad provisioners may, for example, not be as 'uncooperative' as they appear. if they are good defenders. Thus, the distribution of helping tasks between group members should have important implications for our interpretation of group dynamics. Here, we compare two distinct forms of helping behaviour in the cooperatively breeding noisy miner (Manorina melanocephala): chick provisioning and mobbing nest predators. We show that the way in which individual helpers invest in these two helping behaviours varies enormously across individuals and among social groups. Good provisioners often contributed relatively little to mobbing and vice versa. Indeed, (18%) of helpers only mobbed, 22% just provisioned, whereas 60% of helpers performed both forms of helping. Across nests, provisioning was significantly negatively correlated with mobbing effort. We suggest that small differences in the costs or benefits of different aspects of helping (due to differences in age, relatedness or social status) have a big impact on the division of labour within a group. Consequently, social groups can be made up from individuals who often specialise in one helping behaviour, and/or helpers who perform a number of behaviours to differing degrees. Division of labour within social groups will, therefore, have important consequences for the maintenance of cooperatively breeding in vertebrates.

Habitat type and density influence vocal signal design in satin bowerbirds

This study provided a thorough test of the acoustic adaptation hypothesis using a within-species comparison of call structure involving a wide range of habitat types, an objective measure of habitat density and direct measures of habitat-related attenuation. The structure of the bower advertisement call of the satin bowerbird was measured in 16 populations from throughout the species' range and related to the habitat type and density at each site. Transmission of white noise, pure tones and different bowerbird dialects was measured in five of six habitat types inhabited by satin bowerbirds. Bowerbird advertisement call structure converged in similar habitats but diverged among different habitats; this pattern was apparent at both continent-wide and local geographical scales. Bowerbirds' call structures differed with changes in habitat density, consistent with the acoustic adaptation hypothesis. Lower frequencies and less frequency modulation were utilized in denser habitats such as rainforest and higher frequencies and more frequency modulation were used in the more open eucalypt-dominated habitats. The white noise and pure tone transmission measurements indicated that different habitats varied in their sound transmission properties in a manner consistent with the observed variation in satin bowerbird vocalizations. There was no effect of geographical proximity of recording locations, nor was there the predicted inverse relationship between frequency and body size. These findings indicate that the transmission qualities of different habitats have had a major influence on variation in vocal phenotypes in this species. In addition, previously published molecular data for this species suggest that there is no effect of genetic relatedness on call similarity among satin bowerbird populations.

Costs of learning: the dynamics of mixed-prey exploitation by silver perch, Bidyanus bidyanus (Mitchell, 1838)

Although generalist predators have been reported to forage less efficiently than specialists, there is little information on the extent to which learning can improve the efficiency of mixed-prey foraging. Repeated exposure of silver perch to mixed prey (pelagic Artemia and benthic Chironomus larvae) led to substantial fluctuations in reward rate over relatively long (20-day) timescales. When perch that were familiar with a single prey type were offered two prey types simultaneously, the rate at which they captured both familiar and unfamiliar prey dropped progressively over succeeding trials. This result was not predicted by simple learning paradigms, but could be explained in terms of an interaction between learning and attention. Between-trial patterns in overall intake were complex and differed between the two prey types, but were unaffected by previous prey specialization. However, patterns of prey priority (i.e. the prey type that was preferred at the start of a trial) did vary with previous prey training. All groups of fish converged on the most profitable prey type (chironomids), but this process took 15-20 trials. In contrast, fish offered a single prey type reached asymptotic intake rates within five trials and retained high capture abilities for at least 5 weeks. Learning and memory allow fish to maximize foraging efficiency on patches of a single prey type. However, when foragers are faced with mixed prey populations, cognitive constraints associated with divided attention may impair efficiency, and this impairment can be exacerbated by experience. (c) 2005 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.