Killer whale predation on marine mammals at Punta Norte,Argentina; food sharing,provisioning and foraging strategy

Summary The social dynamics of killer whales (Orcinus orca) that hunt marine mammals are apparently highly flexible, though strong individual associations do exist. The killer whales at Punta Norte offer an unusually detailed view of association patterns and foraging behaviour, and suggest a pattern of behaviour that optimizes hunting efficiency with exception only to strong associations between some individuals and the provisioning and training of offspring. The main points from this paper are as follows: First, hunting effort was concentrated where the capture rate was greatest. All pods selectively attacked the prey type for which they had the highest capture rate. The amount of southern sea lion prey captured was approximately equal to the estimated minimum energetic requirement for killer whales based on weight. Secondly, one whale in each pod did the majority of the hunting, and then provisioned the others in the pod. It was clear on numerous occasions that food was shared. A review of reported incidences of killer wales taking marine mammal prey suggests that it is common for a subset of the individuals in a pod to hunt. These results are discussed in the context of the evolution of foraging behaviour.Offprint requests to: A.R. Hoelzel at the first address     

Large carnivore hunting and the social license to hunt

The social license to operate framework considers how society grants or withholds informal permission for resource extractors to exploit publicly owned resources. We developed a modified model, which we refer to as the social license to hunt (SLH). In it we similarly consider hunters as operators, given that wildlife are legally considered public resources in North America and Europe. We applied the SLH model to examine the controversial hunting of large carnivores, which are frequently killed for trophies. Killing for trophies is widespread, but undertaken by a minority of hunters, and can pose threats to the SLH for trophy-seeking carnivore hunters and potentially beyond. Societal opposition to large carnivore hunting relates not only to conservation concerns but also to misalignment between killing for trophies and dominant public values and attitudes concerning the treatment of animals. We summarized cases related to the killing of grizzly bears (Ursus arctos), wolves (Canis lupus), and other large carnivores in Canada, the United States, and Europe to illustrate how opposition to large carnivore hunting, now expressed primarily on social media, can exert rapid and significant pressure on policy makers and politicians. Evidence of the potential for transformative change to wildlife management and conservation includes proposed and realized changes to legislation, business practice, and wildlife policy, including the banning of some large carnivore hunts. Given that policy is ultimately shaped by societal values and attitudes, research gaps include developing increased insight into public support of various hunting policies beyond that derived from monitoring of social media and public polling. Informed by increased evidence, the SLH model can provide a conceptual foundation for predicting the likelihood of transient versus enduring changes to wildlife conservation policy and practice for a wide variety of taxa and contexts.     

Cooperative hunting in lions: the role of the individual

Summary Individually identified lions (Panthera leo) were observed on the open, semi-arid plains in Namibia. Data from 486 coordinated group hunts were analysed to assess cooperation and individual variation in hunting tactics. Group hunts generally involved a formation whereby some lionesses (wings) circled prey while others (centres) waited for prey to move towards them. Those lionesses that occupied wing stalking roles frequently initiated an attack on the prey, while lionesses in centre roles moved relatively small distances and most often captured prey in flight from other lionesses. Each lioness in a given pride repeatedly occupied the same position in a hunting formation. Hunts where most lionesses present occupied their preferred positions had a high probability of success. Individual hunting behaviour was not inflexible, however, but varied according to different group compositions and to variations in the behaviour of other individuals present. The role of cooperative hunting and its apparent advantages within the semi-arid environment of Etosha National Park, Namibia, are discussed.     

The meat-scrap hypothesis: small quantities of meat may promote cooperative hunting in wild chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

A common explanation for hunting in groups is that doing so yields a greater per capita caloric benefit than hunting solitarily. This is logical for social carnivores, which rely exclusively on meat for energy, but arguably not for omnivores, which obtain calories from either plant or animal matter. The common chimpanzee, Pan troglodytes, is one of the few true omnivores that regularly hunts in groups. Studies to date have yielded conflicting data regarding the payoffs of group hunting in chimpanzees. Here, we interpret chimpanzee hunting patterns using a new approach. In contrast to the classical assumption that hunting with others maximizes per capita caloric intake, we propose that group hunting is favored because it maximizes an individual’s likelihood of obtaining important micronutrients that may be found in small quantities of meat. We describe a mathematical model demonstrating that group hunting may evolve when individuals can obtain micronutrients more frequently by hunting in groups than by hunting solitarily, provided that group size is below a certain threshold. Twenty five years of data from Gombe National Park, Tanzania are consistent with this prediction. We propose that our ‘meat-scrap’ hypothesis is a unifying approach that may explain group hunting by chimpanzees and other social omnivores.     

Prey detection in trawling insectivorous bats: duckweed affects hunting behaviour in Daubenton's bat, Myotis daubentonii

Daubenton's bat, a trawling vespertilionid bat species, hunts for insects that fly close to, or rest on, the water surface. During summer, many ponds at which Daubenton's bats hunt become gradually covered with duckweed. The purpose of this study was to investigate the effects of duckweed cover on the hunting behaviour of Daubenton's bats and on the ultrasound-reflecting properties of the water surface. Our study revealed the following. (1) Daubenton's bat avoids water surfaces covered with duckweed. (2) Prey abundance was related to the number of foraging Daubenton's bats but was independent of duckweed cover. (3) When mealworms were presented among standardized amounts of duckweed to naturally foraging Daubenton's bats, they caught significantly less mealworms when the duckweed cover was increased. (4) Measurements with ultrasonic signals show that a water surface covered with duckweed returns a much stronger background echo at small angles (i.e. parallel to the water surface) compared to an uncovered water surface. It seems likely that a cover of duckweed on the water surface interferes with prey detection by masking the echoes returning from prey. (5) It was relatively difficult for the bats to discriminate small patches of duckweed from mealworms. The proposed discrimination mechanism for this trawling bat species suggests that single duckweed patches can also be mistaken for natural prey by Daubenton's bats. Received: 4 January 1998 / Accepted after revision: 19 July 1998     

Shark personalities? Repeatability of social network traits in a widely distributed predatory fish

Interest in animal personalities has generated a burgeoning literature on repeatability in individual traits such as boldness or exploration through time or across different contexts. Yet, repeatability can be influenced by the interactive social strategies of individuals, for example, consistent inter-individual variation in aggression is well documented. Previous work has largely focused on the social aspects of repeatability in animal behaviour by testing individuals in dyadic pairings. Under natural conditions, individuals interact in a heterogeneous polyadic network. However, the extent to which there is repeatability of social traits at this higher order network level remains unknown. Here, we provide the first empirical evidence of consistent and repeatable animal social networks. Using a model species of shark, a taxonomic group in which repeatability in behaviour has yet to be described, we repeatedly quantified the social networks of ten independent shark groups across different habitats, testing repeatability in individual network position under changing environments. To understand better the mechanisms behind repeatable social behaviour, we also explored the coupling between individual preferences for specific group sizes and social network position. We quantify repeatability in sharks by demonstrating that despite changes in aggregation measured at the group level, the social network position of individuals is consistent across treatments. Group size preferences were found to influence the social network position of individuals in small groups but less so for larger groups suggesting network structure, and thus, repeatability was driven by social preference over aggregation tendency.     

Feeding and non-feeding aggression can be induced in invasive shore crabs by altering food distribution

Human activity can have a large impact on surrounding ecosystems. For example, humans alter resource distributions for other species, potentially modifying these species competitive dynamics. These changes in local competitive processes are frequently associated with species invasions. Here, we investigate how differences in resource distribution affect competitive behaviour using the highly invasive European shore crab (Carcinus maenas). Using a controlled laboratory experiment in combination with behaviour assays and social network analysis, we show that individuals feeding in habitats with clumped food distributions are more aggressive than individuals feeding in habitats where food is evenly dispersed, and this aggression is present even on days where crabs are not feeding. Additionally, this persistent aggression can be induced, suggesting that individuals of this invasive species possess the flexibility to modify their competitive behaviours in response to differences in food distributions. Furthermore, we show how these individual responses can lead to changes in overall organisation of aggressive interactions within a population. We discuss these results in relation to how human impacts can have long-term effects on competitive behavioural strategies, and how behavioural flexibility can allow invasive species to colonise and persist in highly impacted sites such as urban ecosystems.     

Toward a new understanding of the links between poverty and illegal wildlife hunting

Conservation organizations have increasingly raised concerns about escalating rates of illegal hunting and trade in wildlife. Previous studies have concluded that people hunt illegally because they are financially poor or lack alternative livelihood strategies. However, there has been little attempt to develop a richer understanding of the motivations behind contemporary illegal wildlife hunting. As a first step, we reviewed the academic and policy literatures on poaching and illegal wildlife use and considered the meanings of poverty and the relative importance of structure and individual agency. We placed motivations for illegal wildlife hunting within the context of the complex history of how wildlife laws were initially designed and enforced to indicate how hunting practices by specific communities were criminalized. We also considered the nature of poverty and the reasons for economic deprivation in particular communities to indicate how particular understandings of poverty as material deprivation ultimately shape approaches to illegal wildlife hunting. We found there is a need for a much better understanding of what poverty is and what motivates people to hunt illegally.     

Animal social networks: an introduction

Network analysis has a long history in the mathematical and social sciences and the aim of this introduction is to provide a brief overview of the potential that it holds for the study of animal behaviour. One of the most attractive features of the network paradigm is that it provides a single conceptual framework with which we can study the social organisation of animals at all levels (individual, dyad, group, population) and for all types of interaction (aggressive, cooperative, sexual etc.). Graphical tools allow a visual inspection of networks which often helps inspire ideas for testable hypotheses. Network analysis itself provides a multitude of novel statistical tools that can be used to characterise social patterns in animal populations. Among the important insights that networks have facilitated is that indirect social connections matter. Interactions between individuals generate a social environment at the population level which in turn selects for behavioural strategies at the individual level. A social network is often a perfect means by which to represent heterogeneous relationships in a population. Probing the biological drivers for these heterogeneities, often as a function of time, forms the basis of many of the current uses of network analysis in the behavioural sciences. This special issue on social networks brings together a diverse group of practitioners whose study systems range from social insects over reptiles to birds, cetaceans, ungulates and primates in order to illustrate the wide-ranging applications of network analysis. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).     

Cooperative prey capture by young subsocial spiders

Matriphagous young of a subsocial spider Amaurobius ferox exhibit collective predation during their post-maternal social period. In this paper, we examine functional mechanisms of collective predation by sibling groups. Predation efficiency increased with increasing number of individuals within each group. Solitary or paired individuals were generally unable to capture a 20 mg cricket. In larger groups, more individuals participated and captured the prey more quickly. Some siblings did not take part in paralyzing prey, but later consumed it. The proportion of these profiteers within a group increased with the group size. Presented with prey of different sizes (1, 5, or 40 mg), siblings were most aggressive towards each other when predating on 5 mg prey. Prey of this size could be captured by a single individual and yet were sufficiently large for more than one individual to eat. Siblings were much less aggressive towards one another during the capture of 40 mg prey, which require the assistance of other individuals to capture. By providing the same mass of prey in different numbers of individuals (a single cricket of 40 or 40 mg of first-instar crickets), we tested the influence of cooperation on the post-maternal social period. We found no difference in the development of young during the social period nor the timing of dispersal and the body mass of dispersing individuals. We conclude that the young of this subsocial animal increased predation efficiency by cooperative hunting after the mother's death.     

Production of food-associated calls in wild male chimpanzees is dependent on the composition of the audience

Chimpanzees produce acoustically distinct calls when encountering food. Previous research on a number of species has indicated that food-associated calls are relatively widespread in animal communication, and the production of these calls can be influenced by both ecological and social factors. Here, we investigate the factors influencing the production of food-associated calls in wild chimpanzees and examine whether male chimpanzees produce food-associated calls selectively in the presence of important social partners. Male chimpanzees form stable long-term social relationships with each other, and these social bonds are vital in enabling a range of cooperative activities, such as group hunting and territory defence. Our data show that males were significantly more likely to produce food-associated calls if an important social partner was nearby, regardless of the size of the audience or the presence of oestrus females. Call production was also mediated by the size of the food patch and by whether or not the food could be monopolised. The presence of important social partners explained most of the variation in male calling behaviour, indicating that food-associated calls are socially directed and serve a bonding function.     

Females go where the food is: does the socio-ecological model explain variation in social organisation of solitary foragers?

The socio-ecological model (SEM) links ecological factors with characteristics of social systems and allows predictions about the relationships between resource distribution, type of competition and social organisation. It has been mainly applied to group-living species but ought to explain variation in social organisation of solitary species as well. The aim of this study was to test basic predictions of the SEM in two solitary primates, which differ in two characteristics of female association patterns: (1) spatial ranging and (2) sleeping associations. Beginning in August 2002, we regularly (re-)captured and marked individuals of sympatric populations of Madame Berthe's and grey mouse lemurs (Microcebus berthae, Microcebus murinus) in Kirindy Forest (Madagascar). We recorded data on spatial patterns, feeding and social behaviour by means of direct observation of radio-collared females. The major food sources of M. berthae occurred in small dispersed patches leading to strong within-group scramble competition and over-dispersed females with a low potential for female associations. In contrast, M. murinus additionally used patchily distributed, high-quality (large) resources facilitating within-group contest competition. The combined influence of less strong within-group scramble and contest as well as between-group contest over non-food resources allowed females of this species to cluster in space. Additionally, we experimentally manipulated the spatial distribution of food sources and found that females adjusted their spatial patterns to food resource distribution. Thus, our results support basic predictions of the SEM and demonstrated that it can also explain variation in social organisation of solitary foragers.     

Feeding ecology and evolutionary survival of the living coelacanth Latimeria chalumnae

One concept of evolutionary ecology holds that a living fossil is the result of past evolutionary events, and is adapted to recent selective forces only if they are similar to the selective forces in the past. We describe the present environment of the living coelacanth Latimeria chalumnae Smith, 1939 at Grande Comore, western Indian Ocean and report depth-dependent cave distribution, temperature, salinity and oxygen values which are compared to the fish's distribution and its physiological demands. We studied the activity pattern, feeding behaviour, prey abundance and hunting success to evaluate possible links between environmental conditions, feeding ecology and evolutionary success of this ancient fish. Transmitter tracking experiments indicate nocturnal activity of the piscivorous predator which hunts between approximately 200 m below the surface to 500 m depth. Fish and prey density were measured between 200 and 400 m, both increase with depth. Feeding tracks and feeding strikes of the coelacanth at various depths were simulated with the help of video and laser techniques. Along a 9447 m video transect a total of 31 potential feeding strikes occurred. Assuming 100% hunting success, medium-sized individuals would obtain 122 g and large females 299 g of prey. Estimates of metabolic rates revealed for females 3.7 ml O₂ kg⁻¹ h⁻¹ and for males 4.5 ml O₂ kg⁻¹ h⁻¹. Today coelacanths are considered to be a specialist deep-water form and to inhabit, with their ancient morphology, a contemporary environment where they compete with advanced, modern fish. Received: 5 July 1999 / Accepted: 11 November 1999     

Flexible bat echolocation: the influence of individual,habitat and conspecifics on sonar signal design

Acoustic signals which are used in animal communication must carry a variety of information and are therefore highly flexible. Echolocation has probably such functions and could prove as flexible. Measurable variabitlity can indicate flexibility in a behaviour. To quantify variability in bat sonar and relate to behavioural and environmental factors, I recorded echolocation calls of Euderma maculatum, Eptesicus fuscus, Lasiurus borealis and L. cinereus while the bats hunted in their natural habitat. I analysed 3390 search phase calls emitted by 16 known and 16 unknown individuals foraging in different environmental and behvioural situations. All four species used mainly multiharmonic signals that showed considerable intra- and inter-individual variability in the five signal variables I analysed (call duration, call interval, highest and lowest frequency and frequency with maximum energy) and also in the shape of the sonagram. A nested multivariate analysis of variance identified the influences of individual, hunting site, close conspecifics and of each observation on the frequency with maximum energy in the calls, and on other variables measured. Individual bats differed in multiple comparisons, most often in the main call frequency and least often in call interval. In a discriminant function analysis with resubstitution, 56–76% of a species' calls were assigned to the correct individual. Distinct individual call patterns were recorded in special situations in all species and the size of foraging areas in forested areas influenced temporal and spectral call structure. Echolocation behaviour was influenced by the presence of conspecifics. When bats were hunting together, call duration decreased and call interval increased in all species, but spectral effects were less pronounced. The role of morphometric differences as the source of individually distinct vocalizations is discussed. I also examined signal adaptations to long range echolocation and the influence of obstacle distance on echolocation call design. My results allow to discuss the problems of echo recognition and jamming avoidance in vespertilionid bats.     

Hunter Reporting of Catch per Unit Effort as a Monitoring Tool in a Bushmeat‐Harvesting System

Abstract: Growing threats to biodiversity in the tropics mean there is an increasing need for effective monitoring that balances scientific rigor with practical feasibility. Alternatives to professional techniques are emerging that are based on the involvement of local people. Such locally based monitoring methods may be more sustainable over time, allow greater spatial coverage and quicker management decisions, lead to increased compliance, and help encourage attitude shifts toward more environmentally sustainable practices. Nevertheless, few studies have yet compared the findings or cost‐effectiveness of locally based methods with professional techniques or investigated the power of locally based methods to detect trends. We gathered data on bushmeat‐hunting catch and effort using a professional technique (accompanying hunters on hunting trips) and two locally based methods in which data were collected by hunters (hunting camp diaries and weekly hunter interviews) in a 15‐month study in Equatorial Guinea. Catch and effort results from locally based methods were strongly correlated with those of the professional technique and the spatial locations of hunting trips reported in the locally based methods accurately reflected those recorded with the professional technique. We used power simulations of catch and effort data to show that locally based methods can reliably detect meaningful levels of change (20% change with 80% power at significance level [α]= 0.05) in multispecies catch per unit effort. Locally based methods were the most cost‐effective for monitoring. Hunter interviews collected catch and effort data on 240% more hunts per person hour and 94% more hunts per unit cost, spent on monitoring, than the professional technique. Our results suggest that locally based monitoring can offer an accurate, cost‐effective, and sufficiently powerful method to monitor the status of natural resources. To establish such a system in Equatorial Guinea, the current lack of national and local capacity for monitoring and management must be addressed.     

Effects of Landscape Features on the Distribution and Sustainability of Ungulate Hunting in Northern Congo

Abstract: Understanding the spatial dimensions of hunting and prey population dynamics is important in order to estimate the sustainability of hunting in tropical forests. We investigated how hunting offtake of vertebrates differed in mixed forest and monodominant forest (composed of Gilbertiodendron dewevrei) and over different spatial extents within the hunting catchment around the logging town of Kabo, Congo. In 9 months of recall surveys with hunters, we gathered information on over 1500 hunting trips in which ungulates were 65% of the species killed and 82% of harvested biomass. Hunters supplied information on animals killed and the hunting trip, including the area visited (i.e., hunting zone; 11 separate zones within a 506 km² catchment or commonly hunted area). Over 65% of all animals were killed in monodominant forest, which made up 28% of the hunting catchment, and zones with small amounts of monodominant forest were used most frequently by hunters. Given the large offtakes from monodominant forests, we suggest that animal dispersal may be maintaining high, localized harvests in these areas. We believe hunters preferred to hunt in monodominant forest because the understory was accessible and that areas with small amounts of monodominant forest and large amounts of mixed forest were more productive. The variation in hunting pressure we found between and within hunting zones differs from past examinations of spatial variation in hunting offtake, where entire hunting catchments were considered population sinks and areas with low to no hunting (no‐take zones) were outside hunting catchments. Future use of no‐take zones to manage hunting should incorporate variability in offtake within hunting catchments.     

Anti-predation behaviour of red colobus monkeys in the presence of chimpanzees

Predator-prey interactions are usually regarded as evolutionary “arms races”, but evidence is still scarce. We examined whether the anti-predation strategies of red colobus monkeys (Procolobus badius) are adapted to the hunting strategies of chimpanzees (Pan troglodytes) in the Taï National Park, Ivory Coast. Taï chimpanzees search for red colobus groups, approach them silently and hunt co-operatively. Our playback experiments and observations of natural encounters revealed that red colobus hid higher up the trees in positions where exposure to the forest floor is minimal and became silent, when chimpanzees were close. They moved away silently through the canopy, when chimpanzees were still at some distance. However, if a group of diana monkeys was nearby in the latter situation, red colobus sought their presence even if they had to move towards the chimpanzees. Chimpanzees refrained from hunting associated red colobus groups, probably because diana monkeys are excellent sentinels for predators approaching over the forest floor. Thus several elements of both the predator's and the prey's strategies correspond to each other. Finally, we compared the interactions between the two species in Taï and in Gombe, Tanzania. We suggest that the difference in size ratio between the two species at the two sites and adaptation of hunting techniques and of escape modes to different forest structures can explain why Gombe red colobus attack chimpanzees while Taï red colobus try to escape. We conclude that predator-prey interactions can indeed lead to evolutionary arms races, with the specific form of co-adaptations depending on environmental factors.     

Social network theory in the behavioural sciences: potential applications

Social network theory has made major contributions to our understanding of human social organisation but has found relatively little application in the field of animal behaviour. In this review, we identify several broad research areas where the networks approach could greatly enhance our understanding of social patterns and processes in animals. The network theory provides a quantitative framework that can be used to characterise social structure both at the level of the individual and the population. These novel quantitative variables may provide a new tool in addressing key questions in behavioural ecology particularly in relation to the evolution of social organisation and the impact of social structure on evolutionary processes. For example, network measures could be used to compare social networks of different species or populations making full use of the comparative approach. However, the networks approach can in principle go beyond identifying structural patterns and also can help with the understanding of processes within animal populations such as disease transmission and information transfer. Finally, understanding the pattern of interactions in the network (i.e. who is connected to whom) can also shed some light on the evolution of behavioural strategies.     

Spatial and temporal heterogeneities in the contact behaviour of rabbits

It is becoming increasingly evident that the social behaviour of many group-living species is more complex than previously assumed and that free mixing of individuals, even within social groups, is rare. This has important implications for ecological processes, such as disease transmission, which are dependent on interactions between individuals. European wild rabbits (Oryctolagus cuniculus) have been considered traditionally as highly sociable animals that mix freely within groups but interact less frequently between groups. We deployed proximity logging devices to quantify the intra- and inter-group contact behaviour of free-living wild rabbits in two populations in a temperate region of Australia. Altogether, 126 rabbits were fitted with proximity loggers at least once during the study. Radio-tracking was carried out alongside proximity data collection to determine the space use of rabbit social groups within the study sites. On average, a rabbit made only 1.54 ± 0.23 (SE) (median = 0.54) contacts per day with each other rabbit carrying a proximity logger in its social group, and the mean daily contact duration was 202 ± 38 s/day (SE) (median = 29 s). Despite the high degree of home range overlap between the neighbouring social groups, inter-group contacts were highly infrequent and brief. Our results demonstrated considerable spatial and temporal heterogeneities in the contact behaviour between individual rabbits, both between populations and between and within social groups in the same population. Such variations in the social organisation of rabbits are likely to create complex patterns of disease transmission through direct contact and may contribute towards observed heterogeneities in the effects of disease on wild rabbit populations.     

Effects of Multiple Levels of Social Organization on Survival and Abundance

Abstract: Identifying how social organization shapes individual behavior, survival, and fecundity of animals that live in groups can inform conservation efforts and improve forecasts of population abundance, even when the mechanism responsible for group‐level differences is unknown. We constructed a hierarchical Bayesian model to quantify the relative variability in survival rates among different levels of social organization (matrilines and pods) of an endangered population of killer whales (Orcinus orca). Individual killer whales often participate in group activities such as prey sharing and cooperative hunting. The estimated age‐specific survival probabilities and survivorship curves differed considerably among pods and to a lesser extent among matrilines (within pods). Across all pods, males had lower life expectancy than females. Differences in survival between pods may be caused by a combination of factors that vary across the population's range, including reduced prey availability, contaminants in prey, and human activity. Our modeling approach could be applied to demographic rates for other species and for parameters other than survival, including reproduction, prey selection, movement, and detection probabilities.