Social dominance modifies behavioral rhythm in a queenless ant

Social insects provide an intriguing model system in chronobiology. Typically, an egg-laying queen exhibits arrhythmicity in activity while foraging worker has clear rhythmicity. In the queenless ant, Diacamma sp., from Japan, colony members lack morphological caste, and reproductive differentiation occurs as a consequence of dominance hierarchy formation. Their specialized dominance interaction “gemmae mutilation”, provide us a fascinating model system to investigate the effect of social dominance on rhythmic ontogeny. Measurement of individual rhythms revealed that they have clear circadian rhythm at eclosion but it is diminished by social mutilation of gemmae. Moreover, unlike highly eusocial species, mated egg-layer (i.e., gamergate) possessed a circadian rhythm even after mating in Diacamma. Measurement of colony-level rhythms revealed that gemmae mutilations are performed in the limited time of the day, but foraging occurs around-the-clock. The above finding is a novel form of temporal organization in social insects, providing a new insight in morphologically casteless species. We discuss the causes and consequences of rhythmic variability in social organization.     

Caste polyethism and collective defense in the ant,Pbeidole pallidula: the outcome of quantitative differences in recruitment

Summary During agonistic encounters, both minors and majors of the European ant P. pallidula actively cooperate in defense. Minors seize the legs of the intruder and in some cases induce the recruitment of nestmates whereas majors kill the spreadeagled alien ant. The defensive strategy of P. pallidula is very flexible and adapted to both the number of alient ants and to the intruder's superiority in fighting. On the one hand, only a massive invasion of alien minors results in a slow mobilization of resident ants to the combat area, elicited by recruiters performing weak tactile invitations and trail-laying behavior. On the other hand, the presence of 10 majors induces a fast and massive recruitment achieved by intense trail-laying and tactile invitations from the recruiters. Because of their high response threshold to invitations, resident majors are mobilized only during these intense recruitments, their exit being additionally enhanced by their preferential stimulation. The adaptiveness of this defensive strategy is discussed. It is also suggested that simple decision-making rules of recruitment and caste differences in behavioral thresholds could account for the complexity of P. pallidula defensive strategies.Senior Research Assistant at Belgian National Fund for Scientific Research Offprint requests to: C. Detrain     

The role of age in temporal polyethism in a primitively eusocial wasp

The relation of age to division of labor was assessed in a primitively eusocial wasp, Ropalidia marginata. The performance of four functionally significant tasks was analyzed. It was found that age has a definite correlation with division of labor, since wasps performed tasks in a distinct sequence in their life with successive tasks being initiated at significantly older ages. Age of a wasp was measured in absolute terms and also relative to other individuals in the colony. Probability of performance of a given task relative to other tasks (PTP) and absolute rates at which tasks were performed per unit time (FTP) both showed clear age-dependent patterns, confirming the association of age with division of labor. The proportion of variance explained for both PTP and FTP was significantly higher with relative age than with absolute age. Interindividual interactions were found to be a potential mechanism through which wasps can determine their relative age. The advantages of work organization depending on relative age and the constraints imposed by absolute age are discussed. Received: 2 April 1997 / Accepted after revision: 20 July 1997     

Genotypic variability in age polyethism and task specialization in the honey bee,Apis mellifera (Hymenoptera: Apidae)

Summary The currently accepted model for division of labor in honey bees, Apis mellifera, explains variation in the frequency at which workers perform specific tasks as the result of differences in age and environment. Although well documented, the model is incomplete because it fails to take genotypic variability among workers into account. We show that workers from two genetically distinct strains of honey bees differed in the age at which they began foraging and in the relative frequency at which they foraged for pollen. Workers from the two strains also exhibited significant spatial heterogeneity within the nest, suggesting that they differed in the frequency at which they performed within-nest tasks as well. A heuristic model of division of labor that incorporates genotypic effects is presented.     

Colony founding,queen dominance and oligogyny in the Australian meat ant Iridomyrmex purpureus

Summary Field observations and laboratory experiments demonstrate that in the Australian meat ant, Iridomyrmex purpureus, the modes of colony founding are remarkably diverse. New colonies can originate from single foundresses (haplometrosis), or foundress associations (pleometrosis), or by colony budding, or the adoption of newly-mated queens that dig founding chambers next to mature nests (probably their natal nests, as workers protect them and may help them dig). Readoption of foundresses and pleometrosis lead to the coexistence of several queens in one nest. We discovered a striking antagonistic behavior among coexisting queens in young colonies, in the form of ritualized antennation bouts. These interactions result in a reproductive rank order in which dominant queens inhibit egg-laying by subordinates, but escalation into physical fighting is rare. Workers ignore queen dominance interactions and treat all queens equally. The first quantitative ethogram of dominance display behavior between multiple ant queens, and its reproductive consequences, is presented. As a colony grows, queens become intolerant of each other's presence and permanently separate within the nest. Once separated, queens appear to be equal in status, laying approximately equal numbers of eggs. All queens continue to be tolerated by workers, even when the colony has reached a size of several thousand workers and begun to produce reproductives. Such mature nests of I. purpureus fulfill the criteria of oligogyny, defined by worker tolerance toward more than one queen and antagonism among queens, such that a limited number of fully functional queens are spaced far apart within a single colony. Oligogynous colonies can arise in this species by pleometrotic founding (primary oligogyny) or by adoption of queens into existing nests (secondary oligogyny). The adaptive significance of the complex system of colony founding, queen dominance and oligogyny in I. purpureus is discussed.     

Increased grooming after repeated brood care provides sanitary benefits in a clonal ant

Repeated pathogen exposure is a common threat in colonies of social insects, posing selection pressures on colony members to respond with improved disease-defense performance. We here tested whether experience gained by repeated tending of low-level fungus-exposed (Metarhizium robertsii) larvae may alter the performance of sanitary brood care in the clonal ant, Platythyrea punctata. We trained ants individually over nine consecutive trials to either sham-treated or fungus-exposed larvae. We then compared the larval grooming behavior of naive and trained ants and measured how effectively they removed infectious fungal conidiospores from the fungus-exposed larvae. We found that the ants changed the duration of larval grooming in response to both, larval treatment and their level of experience: (1) sham-treated larvae received longer grooming than the fungus-exposed larvae and (2) trained ants performed less self-grooming but longer larval grooming than naive ants, which was true for both, ants trained to fungus-exposed and also to sham-treated larvae. Ants that groomed the fungus-exposed larvae for longer periods removed a higher number of fungal conidiospores from the surface of the fungus-exposed larvae. As experienced ants performed longer larval grooming, they were more effective in fungal removal, thus making them better caretakers under pathogen attack of the colony. By studying this clonal ant, we can thus conclude that even in the absence of genetic variation between colony members, differences in experience levels of brood care may affect performance of sanitary brood care in social insects.     

Age-related repertoire expansion and division of labor in Pheidole dentata (Hymenoptera: Formicidae): a new perspective on temporal polyethism and behavioral plasticity in ants

The controversy concerning the extent to which the organization of division of labor in social insects is a developmental process or is based on task allocation dynamics that emerge from colony need independent of worker age and endocrine or neural state has yet to be resolved. We present a novel analysis of temporal polyethism in the ant Pheidole dentata, demonstrating that task attendance by minor workers does not shift among spatially associated sets of behaviors that minimally overlap but rather expands with age. Our results show that the number of tasks performed by older minors increases through the addition and retention of behaviors, with up to a sixfold increase in repertoire size from day 1 to day 20 of adult life. We also show that older minors respond to colony needs by performing significantly more brood care as its demand increases, indicating that they can quickly upregulate nursing according to labor requirements. This level of plasticity was absent in younger siblings. The breadth of responsiveness to task-related olfactory stimuli increased with age. In a binary choice test in which young and old minor workers could orient toward odorants from brood or food, older workers responded to both brood and food, whereas young workers responded only to brood. These dissimilar responses to stimuli associated with nursing and foraging indicate age-related differences in sensory ability and provide a physiological basis for the age-related repertoire expansion model. We discuss repertoire expansion in P. dentata in light of behavioral development and caste flexibility in ants.     

Fire ant thermal preferences: behavioral control of growth and metabolism

Summary Thermal preferences of well-fed and food-limited fire ant colonies (Solenopsis invicta) were studied in relation to colony growth and metabolic costs. The growth curve for well-fed colonies was strongly skewed toward warmer temperatures with maximal growth occurring near 32° C (Fig. 2A). The growth curve for food-limited colonies was skewed toward cooler temperatures with maximal colony size occurring around 25° C (Fig. 2B). Food-limited colonies apparently grew larger at cooler temperatures because metabolic costs of workers were reduced. A series of binary choice tests confirmed three predictions concerning fire ant thermal preferences (Figs. 3–4). First, well-fed colonies preferred brood temperatures very near the optimum for colony growth (31° C versus 32° C). Colonies were also able to select appropriate suboptimal growth temperatures when the optimal range was unavailable. Secondly, as predicted, a large percentage of colony workers ( 30% in well-fed colonies) consistently chose cooler temperatures than those selected for the brood. This strategy probably increases longevity of workers not directly associated with brood care. Thirdly, food-limited colonies preferred cooler temperatures than well-fed colonies. Metabolic costs of food-limited colonies were reduced by approximately 7% because of (1) slightly cooler brood temperatures (30° C versus 31° C) and because (2) an additional 20–30% of the workers selected cooler temperatures. The addition of excess food reversed food-limited thermal preferences within 12 h for the brood (Fig. 5) and several days for the workers. Contrary to expectations, thermal preferences for brood in food-limited colonies did not match the food-limited growth curve, perhaps because fire ant colonies can choose to rear brood at warm temperatures while maintaining accumulated colony biomass at cooler temperatures. Correspondence to: S.D. Porter     

Prey density and the behavioral flexibility of a marine predator: the common murre (Uria aalge)

Flexible time budgets allow individual animals to buffer the effects of variable food availability by allocating more time to foraging when food density decreases. This trait should be especially important for marine predators that forage on patchy and ephemeral food resources. We examined flexible time allocation by a long-lived marine predator, the Common Murre (Uria aalge), using data collected in a five-year study at three colonies in Alaska (USA) with contrasting environmental conditions. Annual hydroacoustic surveys revealed an order-of-magnitude variation in food density among the 15 colony-years of study. We used data on parental time budgets and local prey density to test predictions from two hypotheses: Hypothesis A, the colony attendance of seabirds varies nonlinearly with food density; and Hypothesis B, flexible time allocation of parent murres buffers chicks against variable food availability. Hypothesis A was supported; colony attendance by murres was positively correlated with food over a limited range of poor-to-moderate food densities, but independent of food over a broader range of higher densities. This is the first empirical evidence for a nonlinear response of a marine predator's time budget to changes in prey density. Predictions from Hypothesis B were largely supported: (1) chick-feeding rates were fairly constant over a wide range of densities and only dropped below 3.5 meals per day at the low end of prey density, and (2) there was a nonlinear relationship between chick-feeding rates and time spent at the colony, with chick-feeding rates only declining after time at the colony by the nonbrooding parent was reduced to a minimum. The ability of parents to adjust their foraging time by more than 2 h/d explains why they were able to maintain chick-feeding rates of more than 3.5 meals/d across a 10-fold range in local food density.     

Waste management in the leaf-cutting ant Acromyrmex echinatior: the role of worker size, age and plasticity

Division of labour is the hallmark of the success of many social animals. It may be especially important with regard to waste management because waste often contains pathogens or hazardous toxins and worker specialisation can reduce the number of group members exposed to it. Here we examine waste management in a fungus-farming, leaf-cutting ant, Acromyrmex echinatior, in which waste management is necessary to protect their vulnerable fungal crop. By marking ants with task-specific paint colours, we found clear division of labour between workers that engage in waste management and those that forage, at least during the fine timescale of the 3-day marking period. This division of labour was influenced by both age and size, with waste management workers tending to be smaller and younger than foragers. The role of preventing contaminated ants from entering the colony was fulfilled mainly by medium-sized workers. When the level of waste was experimentally increased, most of the ants that responded to remove the waste were workers previously engaged in tasks inside the nest rather than external waste workers or foragers. These responding workers tended to be young and medium-sized. Surprisingly, the responding ants were subsequently able to revert back to working within the fungus garden, but the probability of them doing so depended on their age and the length of time they were exposed to waste. The results demonstrate the importance of division of labour with regard to waste management in A. echinatior and show that this is adaptable to changing needs.     

Dominance and polyethism in the eusocial wasp Mischocyttarus mastigophorus (Hymenoptera: Vespidae)

Dominance interactions affected patterns of non-reproductive division of labor (polyethism) in the eusocial wasp Mischocyttarus mastigophorus. Socially dominant individuals foraged for food (nectar and insect prey) at lower rates than subordinate individuals. In contrast, dominant wasps performed most of the foraging for the wood pulp used in nest construction. Social dominance also affected partitioning of materials collected by foragers when they returned to the nest. Wood pulp loads were never shared with nest mates, while food loads, especially insect prey, were often partitioned with other wasps. Dominant individuals on the nest were more likely to take food from arriving foragers than subordinate individuals. The role of dominance interactions in regulating polyethism has evolved in the eusocial paper wasps (Polistinae). Both specialization by foragers and task partitioning have increased from basal genera (independent-founding wasps, including Mischo-cyttarus spp.) to more derived genera (swarm-founding Epiponini). Dominance interactions do not regulate forager specialization or task partitioning in epiponines. I hypothesize that these changes in polyethism were enabled by the evolution of increased colony size in the Epiponini. Received: 8 December 1997 / Accepted after revision: 28 March 1998     

Mating frequency, colony size, polyethism and sex ratio in fungus-growing ants (Attini)

The mating frequency of queens was estimated for eight attine ant species, Myrmicocrypta ednaella, Apterostigma mayri, Cyphomyrmex costatus, C. rimosus (four lower attines), Trachymyrmex isthmicus, Serico-myrmex amabalis, Acromyrmex octospinosus and Atta colombica (four higher attines), and correlated to colony size, worker polyethism, and sex ratio. Mating frequency was calculated from within-colony relatedness estimated by CAP-PCR DNA fingerprinting. Most queens of lower attines and T. isthmicus mated with only one male, while those of the three higher attines mated with multiple males. Mating frequency was positively correlated with colony size. Polyethism among workers was dependent on worker age in lower attines but on body size in higher attines, suggesting some correlation between mating frequency (i.e., within-colony gene diversity) and caste complexity. The sex ratio was biased toward females in species where the mating frequency equaled one, but toward males in species where the mating frequency was greater than two. Changing in nest site from ground surface to deep underground may have facilitated the evolution of large colony size in Attini, and this may have resulted in the evolution of polyandry (a queen mates with multiple males). With the evolution of polyandry in higher attines, Atta and Acromyrmex in particular have generated high genetic diversity within their colonies and complex social structures. Received: 26 October 1999 / Revised: 25 May 2000 / Accepted: 24 June 2000     

Inbreeding in a lek-mating ant species, Pogonomyrmex occidentalis

In this paper we have two goals. First, we examine the effects of sample size on the statistical power to detect a given amount of inbreeding in social insect populations. The statistical power to detect a given level of inbreeding is largely a function of the number of colonies sampled. We explore two sampling schemes, one in which a single individual per colony is sampled for different sample sizes and a second sampling scheme in which constant sampling effort is maintained (the product of the number of colonies and the number of workers per colony is constant). We find that adding additional workers to a sample from a colony makes it easier to detect inbreeding in samples from given number of colonies; however, adding more colonies rather than more workers per colony always gives greater power to detect inbreeding. Because even relatively large amounts of sib-mating generate relatively small inbreeding coefficients, detection of even substantial deviations from random mating will require very large samples. Second, we look at the amount of inbreeding in a large population of the western harvest ant, Pogonomyrmex occidentalis. We find deviations from Hardy-Weinberg equilibrium equivalent to approximately 27% sib-mating in our population ( f = 0.09). Review of past studies on the population structure of other Pogonomyrmex species suggests that inbreeding may be a regular feature of the mating system of these ants. Although P. occidentalisis a swarm-mating species, there are a number of features of its population biology which suggest that the effective population size may be small. These include topographical variation that potentially breaks the population into demes, variation in the reproductive output of colonies, and variation in the size of reproductives produced by colonies. Received: 6 May 1996 / Accepted after revision: 6 October 1996     

Social dominance,seasonal movements,and spatial segregation in African elephants: a contribution to conservation behavior

The structure of dominance relationships among individuals in a population is known to influence their fitness, access to resources, risk of predation, and even energy budgets. Recent advances in global positioning system radio telemetry provide data to evaluate the influence of social relationships on population spatial structure and ranging tactics. Using current models of socio-ecology as a framework, we explore the spatial behaviors relating to the maintenance of transitive (i.e., linear) dominance hierarchies between elephant social groups despite the infrequent occurrence of contests over resources and lack of territorial behavior. Data collected from seven families of different rank demonstrate that dominant groups disproportionately use preferred habitats, limit their exposure to predation/conflict with humans by avoiding unprotected areas, and expend less energy than subordinate groups during the dry season. Hence, our data provide strong evidence of rank derived spatial partitioning in this migratory species. These behaviors, however, were not found during the wet season, indicating that spatial segregation of elephants is related to resource availability. Our results indicate the importance of protecting preexisting social mechanisms for mitigating the ecological impacts of high density in this species. This analysis provides an exemplar of how behavioral research in a socio-ecological framework can serve to identify factors salient to the persistence and management of at risk species or populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Chemical composition of Eurasian lynx urine conveys information on reproductive state,individual identity,and urine age

In mammals, the chemical profiles of individuals are complex and variable mixtures, and animals perceive information based on variation in the overall quality of these mixtures. A variety of compounds potentially involved in chemical communication have been characterized in the urine of different felid species, but little is known about the information content of felid scent marks. In this study, we investigated whether chemical composition of Eurasian lynx Lynx lynx urine was related to sex, reproductive state, and individual identity. We further analysed if elemental sulphur in lynx urine could serve as a dietary cue or as an indicator for the freshness of a scent mark. We collected urine from captive and wild Eurasian lynx, and analysed volatile constituents of urine by means of solid phase microextraction and gas chromatography-mass spectrometry. Our results show that lynx scent profiles contain sex-specific information on reproductive state, as well as individual identity cues. Urine marks are, therefore, well-suited to fulfil a role in reproductive behaviour and social organisation of wild lynx populations. Relative sulphur content was unrelated to time since last feeding, but decreased with age of the urine sample. The influence of diet and body condition on scent profiles should be further investigated by means of experimental studies, and may shed more light on the messages encoded in carnivore scent-marks.     

Hierarchical dominance structure in reintroduced California condors: correlates, consequences, and dynamics

Populations of reintroduced California condors (Gymnogyps californianus) develop complex social structures and dynamics to maintain stable group cohesion, and birds that do not successfully integrate into group hierarchies have highly impaired survivability. Consequently, improved understanding of condor socioecology is needed to inform conservation management strategies. We report on the dominance structure of free-ranging condors and identify the causes and consequences of rank in condor populations by matching social status with the behavioral and physical correlates of individual birds. We characterized the hierarchical social structure of wild condor populations as mildly linear, despotic, and dynamic. Condor social groups were not egalitarian and dominance hierarchies regulated competitive access to food resources. Absence of kin-based social groups also indicated that condor social structure is individualistic. Agonistic interactions among condors were strongly unidirectional, but the overall linearity and steepness of their hierarchies was low. Although one aggressive male maintained the highest dominance rank across the 3-year observation period, there was considerable fluidity in social status among condors within middle and lower rank orders. Older condors were more dominant than younger birds and younger males supplanted older females over time to achieve higher status. Dominance rank did not predict the amount of time that a bird spent feeding at a carcass or the frequency that a bird was interrupted while feeding. Thus, younger, less dominant birds are able to obtain sufficient nutrition in wild social populations.