Vibrational communication facilitates cooperative foraging in a phloem-feeding insect

Insects are the dominant herbivores in tropical forests, with a range of mechanisms for exploiting plant resources. For group-living species, such mechanisms may involve communication. The Neotropical treehopper Calloconophora pinguis (Hemiptera: Membracidae) is a sap-feeding species in which groups of siblings feed on new leaves during the brief period of leaf expansion. Using an experimental approach, a process of cooperative foraging among siblings was documented, in which a few individuals in a group behave as scouts, locating a new feeding site and advertising it using plant-borne vibrational signals. Signalling leads to a period of positive feedback in which newly recruited individuals signal in concert with those already there. The food signalling system of C. pinguis is unique in its use of synchronized group displays and in the tight coordination of receiver responses with collective signals. Examples from a number of taxonomic groups show that vibrational communication can allow group-living insects to solve the challenges of feeding on plants, such as remaining in a foraging group or avoiding predation. While most research has focused on leaf-feeding species, sap-feeding species may remove just as much biomass. This study shows that cooperative vibrational communication underlies the ability of a sap-feeding species to exploit plant resources during a narrow window of availability.     

Do deposed adult male Sichuan snub-nosed monkeys Rhinopithecus roxellana roam as solitary bachelors or continue to interact with former band members?

Dispersal patterns exhibited by group-living animals play an important role in individual reproductive success and survival,as well as population and social dynamics(Greenwood,1980;Isbell and van Vuren,1996;Kappeler and van Schaik,2002).Dispersing individuals may emigrate voluntarily,or they may be forcibly evicted by other individuals(Pusey and Packer,1987;Clobert et al.,2001).One cause of individual dispersal is when an extra-group male compels another male to leave a group of females.This phenomenon has ...     

Spacing pattern in a social group of stray cats: effects on male reproductive success

Reproductive consequences of male spacing patterns have received relatively little attention in nonterritorial mammals, in particular in group-living species, where most studies have focused on the relation between social rank and reproductive success. We investigated the effects of spacing pattern on male reproductive success within a social, nonterritorial, promiscuous population of stray cats, Felis catus. Male home ranges overlapped home ranges of many females, consistent with a promiscuous mating system. Furthermore, males with the largest home ranges included the most female home ranges; they successfully reproduced with these females and had the highest reproductive success. Home range size predicted male reproductive success even when controlling for the effect of social rank. However, males also reproduced with females whose home range did not overlap their home range, suggesting that males can make quick excursions outside their home range to find new mating opportunities. We conclude that, in group-living situations, a male's ability to maintain a large home range may be one of the principal causes of variation in mating success in the stray cat.     

Complexity,dynamics and diversity of sociality in group-living mammals

Numerous studies in group-living animals with stable compositions have demonstrated the complex and dynamic nature of social behaviour. Empirical studies occasionally provide principles that cannot be applied directly to other group-living species. Because of this, researchers are required to address fine-scaled conceptual questions and to incorporate species-specific characteristics of the study species. In this paper, I raise three key topics that will promote our understanding of animal sociality: the effects of heterogeneous social relationships on the pattern, distribution, and function of social interactions; conflict management for maintaining group living; and meta-dyad-level perspectives for understanding dyadic social relationships and behaviours. Through the discussion of these topics together with examples of group-living mammals, I emphasise the importance of direct behavioural observations and functional analyses in studies of species- or taxonomic-group-specific characteristics of social behaviour in a wide range of taxonomic groups. In addition to approaches focusing on specificity, another approach that examines the general principles or common characteristics found across different taxonomic groups could provide synthetic and reductive frameworks to understand divergent sociality. The complementary use of these two approaches will offer a comprehensive understanding of social evolution in group-living animals. Nobuyuki Kutsukake is the recipient of the 12th Denzaburo Miyadi Award.     

Group-living and the richness of the parasite fauna in Canadian freshwater fishes

Summary An increased transmission of ectoparasites among individual animals is considered to be an inevitable cost of living in groups, since several kinds of ectoparasites require close proximity between large numbers of hosts for successful transmission. However, we do not know whether individuals belonging to group-living species incur a greater risk of ectoparasitism than individuals of solitary species. Here, using published data from 3 families (60 species) of Canadian freshwater fishes, I test the hypothesis that group-living species are host to more species of contagious ectoparasites (copepods and monogeneans) than solitary host species. As the different fish species have been studied with varying intensity, I used the mean number of parasite species recorded per study as a standard measure of parasite numbers. Multiple regression analyses were performed separately for each family to determine the effects of group-living and of 3 other variables (host size, age, and range) on the richness of the recorded parasite fauna. Once the effects of the other variables were removed, I found no significant effect of sociality on the richness of the parasite fauna per fish species, for contagious ectoparasites and other types of parasites. Neither of the other variables had any influence on the numbers of parasite species per fish species. These results suggest that a richer ectoparasite fauna is not a cost of group-living in fishes.     

Correlates of Male Consortship Rate in Free-Ranging Rhesus Macaques (Macaca mulatta)

Male–male competition for access to receptive females can take the form of nonrecurring fights and/or a sustained contest over mating opportunities. Male physical condition has been linked to dominance rank and reproductive success in species characterized by intrasexual fights for dominance and access to females. In group-living species characterized by endurance rivalry, however, factors contributing to male reproductive success are less well understood. In such species, particularly seasonally breeding ones with low sexual dimorphism and seniority-based rank, age and social factors other than rank may prove important. In the absence of genetic data, male mate guarding or consortship can serve as an indicator of male reproductive success. To evaluate the contribution of age and intragroup sociality to male consortship rate, I collected behavioral data during one nonmating and one mating season in two social groups of free-ranging rhesus macaques that experience no predation or food scarcity. Higher-ranking males, younger males, or males that exhibited lower rate of intrasexual aggression had higher consortship rates. Male–female dyads that groomed outside consortship did not form consortships as often as dyads that did not engage in nonconsort grooming. This is the first study to identify the significance of male–male aggression and male–female affiliation to male consortship rate in a species characterized by endurance rivalry, high male rank stability, and strong female mate choice. Social behaviors and male age may be particularly important in determining male reproductive success in populations experiencing high food availability and a lack of predation, which are typical of an increasing number of vertebrates in areas densely populated by humans.     

Contrasting effects of living in heterosexual pairs and mixed groups on the structure of social attraction in squirrel monkeys (Saimiri).

The structure of social attraction was assessed in pair- and group-living squirrel monkeys (Saimiri) using paired-comparison and single-stimulus preference tests. Effects of the social environment were most prominent in females. Females housed with a single male showed sharply increased attraction to like-sex strangers and less pronounced increase in attraction to opposite-sex strangers, as compared to group-living females. Differences between pair- and group-living males were in the same direction found with females, but less extreme. Most group-living monkeys strongly preferred familiar animals to strangers regardless of sex, and like-sex to opposite-sex familiars. Change in the structure of social attractions in response to variations in the composition of the social environment appears to be an important factor in the maintenance of the species-typical grouping pattern in the squirrel monkey.     

Kin recognition and incest avoidance in a group-living insect

Mate choice theories predict that animals evolved strategiesto mate with optimally genetically dissimilar partners, providingfitness benefits. In group-living species, when adults do notdisperse, assessment of relatedness between conspecifics canbe a key factor for choosing mates. Here, we report for thefirst time, kin recognition abilities and their implicationin mate choice in the gregarious cockroach, Blattella germanica(L.). Binary choice tests showed that females mated preferentiallywith nonsibling rather than with sibling males, thus avoidingincest. In addition, inbreeding induced an important decreaseof their reproductive success. Contrary to what could be expectedwhen females had the choice between a nonsibling strain memberand a nonstrain member, they did not avoid mating with distantlyrelated nonstrain members, and extreme outbreeding induced anincrease of their reproductive success. Furthermore, our matechoice experiments disentangled the influences of familiarityfrom those of relatedness and evidenced that kin discriminationwas based on genetic cues independently of familiarity. Phenotypematching was a plausible mechanism for kin recognition. Contraryto many insect species, body size was not a salient criterionfor mate choice and had no consequences on reproductive success.     

Do Feeding Resources Induce the Adoption of Resource Defence Polygyny in a Lekking Butterfly?

Among polygynous species, males often compete for the possession of mating sites to increase their reproductive success. Weaker individuals frequently adopt alternative non‐territorial mate‐locating tactics, but the adoption of alternative territorial tactics may also occur. Although alternative tactics with territory defence are less common in arthropods, factors that drive its adoption may provide information to understand the organization of different territorial mating systems in the group. Here we investigate the adoption of resource‐based territoriality as an alternative to a non‐resource‐based one by males of the butterfly Paryphthimoides phronius. Male P. phronius commonly defend sunny clearings lacking feeding resources in the forest edge (non‐resource‐based territoriality). However, after experimentally offering fermenting fruit in previously undefended sites, we showed that males also adopt a resource defence tactic. Males in territories with fermenting fruits apparently feed on this resource when they are not defending the territory. However, males in sites without resources did not migrate to territories with resources when given the opportunity. To our knowledge, this is the first experimental investigation to show a non‐resource‐ and an alternative resource‐based mate‐locating tactic in a butterfly. We suggest that this behavioural flexibility may represent an important step to understand the ecological factors responsible for the organization and evolution of different territorial mating systems in insects.     

Endocrinology of sociality: Comparisons between sociable and solitary individuals within the same population of African striped mice

The social organization of species ranges from solitary-living to complex social groups. While the evolutionary reasons of group-living are well studied, the physiological mechanisms underlying alternative social systems are poorly understood. By studying group-living and solitary individuals of the same species, we can determine hormonal correlates of sociality without the problem of confounding phylogenetic factors. The African striped mouse (Rhabdomys pumilio) is a socially flexible species, which can be solitary or alternatively form complex family groups, depending on population density and the extent of reproductive competition. We predicted group-living striped mice to show signs of reproductive suppression and social stress, resulting in higher corticosterone but lower testosterone levels when compared to solitary-living individuals. To determine whether differences in social organization correlated with hormonal differences, we collected blood samples from free-living striped mice during four breeding seasons when we experimentally induced solitary-living in philopatric individuals by locally reducing population density. Striped mice that were group-living did not change their corticosterone or estosterone levels during the study, indicating that there was no temporal effect during the breeding season. Striped mice of both sexes had significantly lower corticosterone levels after switching from group- to solitary-living. Solitary males – but not solitary females – had higher testosterone levels than group-living conspecifics. Our results suggest that group-living results in physiological stress and can induce reproductive suppression, at least in philopatric males. The switch to solitary-living may thus be a tactic to avoid reproductive competition within groups, and is associated with decreased stress hormone levels and onset of independent reproduction.     

Bullia digitalis (Gastropoda) actively pursues moving prey by swash-riding

An individual's success at locating resource items is largely determined by the manner in which it searches. Studies on the movement trajectories of individuals have therefore been useful in elucidating mechanisms by which animals find their resources. Studies thus far have concentrated on animals that live in physically stable environments and search for stationary food items. Several of these studies revealed a surprising ineptitude of individuals in locating their resources. Our study on the other hand focuses on an animal living in a highly turbulent environment that needs to locate moving food items. We studied Bullia digitalis , a whelk that lives in the surf and swash zones of sandy beaches. In spite of constraints imposed by wave action on its movement and obvious difficulties in using positive chemotaxis in a turbulent environment, B. digitalis appears to be most efficient in tracking its prey through the surf. It uses wave-generated flow in a form of locomotion called swash-riding which is commonly used by a wide variety of sandy beach invertebrates.     

The Evolution of Reliable and Unreliable Badges of Fighting Ability

SYNOPSIS. When a population may be characterized by interferencecompetition for resources, variation in fighting ability amongindividuals, and repeated confrontations between individuals,together with difficulty of individual recognition, badges ofstatus should invade as recognition marks that render good fightersmemorable. Reliability of such badges can be maintained by negativefrequency-dependent selection when individuals of differentappearance (and status) either play mutually beneficial rolesor employ alternate competitive tactics. In territorial socialsystems intraspecific mimicry of recognition badges should evolvebecause, in contrast to group-living situations, the cost toa cheat of being discovered is low when individuals are dispersed.The general result of such mimicry is that good and poor fightersbecome similar in appearance. From the theoretical treatmentof status recognition badges I derive a number of predictionsthat pertain both to interand intraspecific differences in conspicuouscoloration and to the evolution of local song dialects in birds.     

A review of the evolutionary causes of rodent group-living

I analyze and summarize the empirical evidence supporting alternative hypotheses posed to explain the evolution of rodent group-living. Eight hypotheses are considered: two rely on net fitness benefits to individuals, five rely on ecological and life-history constraints, and one uses elements of both. I expose the logic behind each hypothesis, identify its key predictions, examine how the available evidence on rodent socioecology supports or rejects its predictions, and identify some priorities for future research. I show that empirical support for most hypotheses is meager due to a lack of relevant studies. Also, empirical support for a particular hypothesis, when it exists, comes from studies of the same species used to formulate the original hypothesis. Two exceptions are the hypothesis that individual rodents live in groups to reduce their predation risk and the hypothesis that group-living was adopted by individuals to reduce their cost of thermoregulation. Finally, most hypotheses have been examined without regard to competing hypotheses and often in a restricted taxonomic context. This is clearly an unfortunate situation given that most competing hypotheses are not mutually exclusive. I suggest that in the future comparative approaches should be used. These studies should examine simultaneously the relevance of different benefits and constraints hypothesized to explain the evolution of rodent sociality.     

Ecological constraints and benefits of philopatry promote group-living in a social but non-cooperatively breeding fish

Why non-breeding subordinates of many animal societies tolerate group-living remains a pertinent question in evolutionary biology. The ecological constraints and benefits of philopatry hypotheses have the potential to explain the maintenance of group-living by specifying the ecological conditions favouring delayed dispersal over independent breeding by subordinates. In this study, I used field and laboratory experiments to investigate the role of ecological and social factors on the dispersal decisions of non-breeding subordinates in the coral-dwelling fish, Paragobiodon xanthosomus (Gobiidae). Subordinate dispersal was strongly influenced by ecological constraints (habitat saturation and risks of movement) and benefits of philopatry (relative coral size). Social factors, namely social rank and forcible eviction, did not affect the occurrence of subordinate dispersal. These results suggest that selection has favoured subordinate P. xanthosomus, which employ a mixed strategy—switching tactics in response to three ecological factors—despite having low mobility and extreme habitat-specific requirements. Furthermore, this study demonstrates the generality of the ecological constraints and benefits of philopatry hypotheses as explanations for group-living in species where subordinates are unrelated to breeders, provide no help and do not strictly delay dispersal.     

Experimental evidence of environmental effects on age-specific reproductive success: the importance of resource quality.

Age-specific access to high-quality resources (e.g. territory or nest site) might be an important determinant for improved reproductive performance with increasing age. I experimentally investigated the effects of territory quality versus other age-related improvements in breeding competence (e.g. foraging skills, breeding experience and local knowledge) on age-specific reproductive success. Territory quality (i.e. territory field layer height) was manipulated in year 2 of northern wheatears (Oenanthe oenanthe) that were breeding in the same territory in two consecutive years. Changing territory quality by changing field layer height had a strong effect on within-individual change in the reproductive success of wheatears. This effect was mainly due to a corresponding change in nest predation risk. When territory quality was kept constant (i.e. no between-year change in territory field layer height), within-individual reproductive success did not change between subsequent years. Thus, age-related improvements in foraging skills, breeding experience and local familiarity had no significant effect on within-individual changes in reproductive success. Increased reproductive success with increased age in northern wheatears is therefore mainly explained by an improved access to high-quality territories with increasing age. I conclude that age-dependent access to high-quality breeding resources might be a widespread phenomenon in nature.     

The spores of Phytophthora: weapons of the plant destroyer

Members of the genus Phytophthora are among the most serious threats to agriculture and food production, causing devastating diseases in hundreds of plant hosts. These fungus-like eukaryotes, which are taxonomically classified as oomycetes, generate asexual and sexual spores with characteristics that greatly contribute to their pathogenic success. The spores include survival and dispersal structures, and potent infectious propagules capable of actively locating hosts. Genetic tools and genomic resources developed over the past decade are now allowing detailed analysis of these important stages in the Phytophthora life cycle.     

Resource-Ratio Theory and the Control of Invasive Plants

There have been relatively few tests of resource-ratio theory in terrestrial systems. Additionally, resources are known to be an important factor determining the success of invasive species. Here I discuss how the study by Newingham and Belnap (pp. 29–40, this issue) tests predictions of resource-ratio theory and how they apply it to questions of invasion by Bromus tectorum in a terrestrial grassland.     

Selection of line-transect methods for estimating the density of group-living animals: lessons from the primates

We review the four major contemporary methods for estimating density of group-living animals from line-transect sampling: perpendicular modelling of group centers, perpendicular modelling of center of measurable individuals, strip transects and animal-observer distance. The efficacy of each method is evaluated to produce a simple selection guide. We review the literature and use field data from the Udzungwa Mountains, Tanzania. The review is relevant to all group-living animals; however, examples are drawn from the primates. Perpendicular methods have better mathematical justification than non-perpendicular methods. For perpendicular methods using detection function models, it is preferable to measure group location using center of measurable individuals, as group centers are hard to estimate. The assumptions of detection function models are often broken in poor visibility habitats or with unhabituated animals. Alternatively strip transects may be used where there are reliable data on group spread and/or visibility. Strip transects are also the most practical, along with the animal-observer method; however, the latter lacks mathematical justification. We conclude that there are arguments for continued use of all four methods. In certain situations the use of raw encounter rates may also be considered. The appropriate method is determined by minimizing bias and considering time, resources and field conditions.     

The cost of being alone: the fate of floaters in a population of cooperatively breeding pied babblers Turdoides bicolor

The occurrence of group-living behaviour has often been explained by the benefits individuals receive through cooperation; including increased reproductive output, vigilance against predators, and load-lightening behaviour. However, to fully understand the benefits of group-living, it is important to quantify the costs of living alone. Here, we look at the fate of floaters (individuals who have no fixed territory and remain alone for extended periods) in a population of cooperatively breeding pied babblers Turdoides bicolor . We found that individuals spent less time foraging and more time vigilant for predators when found as a floater compared to when they were in a group. Consequently, they suffered a continuous loss of body mass, with long-term floaters suffering the highest losses. This had a long-term effect: floaters that eventually did regain a position in a group usually entered as helpers, in contrast to dispersers, who usually entered a new group as breeders. This high cost of living alone highlights the benefits of group-living and may help to understand patterns of delayed dispersal in some social species.     

Quorum decision-making in foraging fish shoals

Quorum responses provide a means for group-living animals to integrate and filter disparate social information to produce accurate and coherent group decisions. A quorum response may be defined as a steep increase in the probability of group members performing a given behaviour once a threshold minimum number of their group mates already performing that behaviour is exceeded. In a previous study we reported the use of a quorum response in group decision-making of threespine sticklebacks (Gasterosteus aculeatus) under a simulated predation threat. Here we examine the use of quorum responses by shoals of sticklebacks in first locating and then leaving a foraging patch. We show that a quorum rule explains movement decisions by threespine sticklebacks toward and then away from a food patch. Following both to and from a food patch occurred when a threshold number of initiators was exceeded, with the threshold being determined by the group size.