The social network structure of a wild meerkat population: 2. Intragroup interactions

Knowledge of the structure of networks of social interactions is important for understanding the evolution of cooperation, transmission of disease, and patterns of social learning, yet little is known of how environmental, ecological, or behavioural factors relate to such structures within groups. We observed grooming, dominance, and foraging competition interactions in eight groups of wild meerkats (Suricata suricatta) and constructed interaction networks for each behaviour. We investigated relationships between networks for different social interactions and explored how group attributes (size and sex ratio), individual attributes (tenure of dominants), and ecological factors (ectoparasite load) are related to variation in network structure. Network structures varied within a group according to interaction type. Further, network structure varied predictably with group attributes, individual attributes, and ecological factors. Networks became less dense as group size increased suggesting that individuals were limited in their number of partners. Groups with more established dominant females were more egalitarian in their grooming and foraging competition interactions, but more despotic in their dominance interactions. The distribution of individuals receiving grooming became more skewed at higher parasite loads, but more equitable at low parasite loads. We conclude that the pattern of interactions between members of meerkat groups is not consistent between groups but instead depends on general attributes of the group, the influence of specific individuals within the group, and ecological factors acting on group members. We suggest that the variation observed in interaction patterns between members of meerkat groups may have fitness consequences both for individual group members and the group itself.     

The social network structure of a wild meerkat population: 1. Inter-group interactions

Groups of individuals frequently interact with each other, but typically analysis of such interactions is restricted to isolated dyads. Social network analysis (SNA) provides a method of analysing polyadic interactions and is used to analyse interactions between individuals. We use a population of 12 groups (ca. 250 animals) of wild meerkats (Suricata suricatta) to test whether SNA can also be used to describe and elucidate patterns of inter-group interactions. Using data collected over 24 months, we constructed two sets of networks, based on direct encounters between groups and instances of roving males visiting other groups. We analysed replicated networks of each type of interaction to investigate similarities between networks of different social interactions as well as testing their stability over time. The two network types were similar to each other when derived from long-term data, but showed significant differences in structure over shorter timescales where they varied according to seasonal and ecological conditions. Networks for both types of inter-group interaction constructed from data collected over 3 months reliably described long-term (12- and 24-month) patterns of interactions between groups, indicating a stable social structure despite variation in group sizes and sex ratios over time. The centrality of each meerkat group in roving interactions networks was unaffected by the sex ratio of its members, indicating that male meerkats preferentially visit geographically close groups rather than those containing most females. Indeed, the strongest predictors of network structure were spatial factors, suggesting that, in contrast to analyses of intra-group interactions, analyses of inter-group interactions using SNA must take spatial factors into account.     

A model of deforestation by human population interactions

A differential equation was employed in modelling deforestation by human population interactions to yield an explicit mathematical model. The theoretical relation and many possible models were applied to the grid cell data in Hiroshima Prefecture, and relative appropriateness of each model was evaluated by Akaike's information criterion (AIC) using raw data. Intensive further verification was executed bythe bootstrap method. It was demonstrated that the theoretical relation was in the best agreement among many other models in comparison.     

Honeybee recruitment to scented food sources: correlations between in-hive social interactions and foraging decisions

Information exchange of environmental cues facilitates decision-making processes among members of insect societies. In honeybee foraging, it is unknown how the odor cues of a resource are relayed to inactive nest mates to enable resource exploitation at specific scented sources. It is presumed that bees need to follow the dance or to be involved in trophallaxis with a successful forager to obtain the discovered floral scent. With this in mind, we evaluated the influence of food scent relayed through in-hive interactions and the subsequent food choices. Results obtained from five colonies demonstrated that bees arriving at a feeding area preferred to land at a feeder carrying the odor currently exploited by the trained forager. The bees that landed at this feeder also showed more in-hive encounters with the trained forager than the individuals that landed at the alternative scented feeder. The most frequent interactions before landing at the correct feeder were body contacts with the active forager, a behavior that involves neither dance following nor trophallaxis. In addition, a reasonable proportion of successful newcomers showed no conspicuous interactions with the active forager. Results suggest that different sources of information can be integrated inside the hive to establish an odor-rewarded association useful to direct honeybees to a feeding site. For example, simple contacts with foragers or food exchanges with non-active foragers seem to be enough to choose a feeding site that carries the same scent collected by the focal forager.     

Adding complex trophic interactions to a size-spectral plankton model: Emergent diversity patterns and limits on predictability

A new model in the NPZ (nutrient-phytoplankton-zooplankton) style is presented, mechanistically simple but with 40 size classes each of phytoplankton (1-20 μm) and small zooplankton (2.1-460 μm), in order to resolve one level of trophic interactions in detail. General, empirical allometric relationships are used to parameterize both the optimal prey size and size selectivity for each grazer class, as is rarely done. This inclusion of complex predator-prey linkages and realistic prey preferences yields a system with an emergent pattern of phytoplankton diversity consistent with global ocean observations, i.e., a parabolic relationship between diversity (as measured by the Shannon evenness) and biomass. It also yields significant long-term time evolution, which places limits on the extent to which the community response to nutrient forcing can be predicted from forcing in a pragmatic sense. When a simple annual cycle in nutrient supply is repeated exactly for many years, transient fluctuations up to a factor of two in spring bloom magnitude persist for 10-20 years before a stable seasonal biomass cycle is achieved. When the amplitude of the nutrient-supply annual cycle is given a random interannual modulation, these long-lived transients add significant noise to a 100-year correlation between annual-mean nutrient supply and annual-mean biomass. This noise is 20% of total interannual variance in the model base case, and ranges from 0% to 40% depending on the grazer size selectivity. In general, unpredictability on the bloom timescale is damped when food-web complexity is increased by making grazers less selective, while unpredictability on the interannual scale shows the opposite pattern, increasing with increasing food-web complexity up to a high threshhold, past which community structure and biomass time evolution both suddenly simplify. These results suggests a new strategy for ensemble ecosystem forecasting and uncertainty estimation, analogous to methods common in circulation and climate modeling, in which internal variability (predator-prey interactions in the biological case; eddies and climate-system oscillations in the physical case) are resolved and quantified, rather than suppressed.     

The number of males in primate social groups: a comparative test of the socioecological model

As applied to polygynous mammals, the socioecological model assumes that environmental risks and resources determine the spatial and temporal distribution of females, which then sets male strategies for monopolizing fertile matings. The effects of female spatial distribution (i.e., female number) and temporal overlap (female mating synchrony) have been examined in comparative studies of primates, but the relative influence of these two factors on male monopolization potential (the number of males) remains unclear. One particular problem is that female synchrony is more difficult to estimate than female number. This paper uses multivariate statistical methods and three independent estimates of female synchrony to assess the roles of spatial and temporal effects in the context of a phylogenetically corrected dataset. These analyses are based on sensitivity analyses involving a total of four phylogenies, with two sets of branch length estimates for each tree, and one nonphylogenetic analysis in which species values are used (because male behavior may represent a facultative response to the distribution of females). The results show: (1) that breeding seasonality predicts male number (statistically significant in six out of nine sensitivity tests); (2) that expected female overlap, after controlling for female group size using residuals, also accounts for the number of males in primate groups (significant in eight out of nine tests), and (3) that actual estimates of female mating synchrony predict male number, again after correcting for female group size (significant in five out of nine tests). Nonsignificant results are in the predicted direction, and female group size is significant in all statistical tests. These analyses therefore demonstrate an independent influence of female temporal overlap on male monopolization strategies in mammalian social systems. Received: 24 July 1998 / Received in revised form: 5 February 1999 / Accepted: 7 February 1999     

Influence of residency and social odors in interactions between competing native and alien rodents

Residency status of individuals in populations may be an important determinant of the outcomes of interspecific competition between native and introduced species. We examined direct behavioral interactions between two similarly sized rodents, the alien Rattus rattus and native Rattus fuscipes when they were respective residents and intruders in a small enclosure. Resident individuals were dominant in their behaviors toward intruders irrespective of the species that was resident. In contrast, interactive behaviors between conspecifics were often neutral or amicable, supporting suggestions that R. rattus and R. fuscipes are social animals. We then tested whether rodent species use heterospecific odors to avoid aggressive competitive interactions and partition space in the field. Neither R. fuscipes nor R. rattus responded to traps scented with the odors of male or female heterospecifics. If R. fuscipes does not recognize the odor of introduced R. rattus, then odors will not be cues to the presence or territorial space of competing heterospecifics. Rather, findings from both enclosure and field trials suggest that direct aggressive interactions between individual R. rattus and R. fuscipes probably facilitate segregation of space between these two species in wild populations, where resident animals may typically be the winners and exclude heterospecific intruders. These findings have implications for the invasion success of introduced rodents such as R. rattus into intact forests, where native populations may have competitive advantage because of their residency status.     

Aggressive thresholds in Dendropsophus ebraccatus: habituation and sensitization to different call types

Males in many chorusing anuran species use aggressive calls during defense of calling spaces from other males. The minimal intensity of another male’s vocalizations that elicits an aggressive call response has been termed the aggressive threshold. Previous studies of aggressive thresholds have shown that they are plastic: males habituated (increased their aggressive thresholds) in response to repeated presentation of stimuli above initial threshold levels. Habituation likely contributes to the stable chorus structure of these species, in which aggressive calling is rare compared to advertisement calls. I have observed high levels of aggressive calling in the treefrog Dendropsophus ebraccatus, suggesting that males of this species do not habituate. In this study, I investigated the plasticity of aggressive thresholds in D. ebraccatus. I measured the aggressive thresholds of males before and after suprathreshold stimulation by both advertisement and aggressive calls. I found that the different call types had different effects: males habituated to advertisement calls but lowered their aggressive thresholds in response to aggressive calls. I consider the latter response to be an example of sensitization, a behavior that has been documented infrequently in vocalizing anurans. Sensitization is a plausible mechanism responsible for the high levels of aggressive calling observed in this species. Given the high costs of aggressive calling, however, it is unclear why a mechanism that increases aggressive call output would be maintained.     

Active and effective nest defence by males in a social apoid wasp

Altruism in the social Hymenoptera is generally considered to be a feature of females rather than males. A popular explanation for this is that in the solitary ancestors of today's social species, males provided little brood care. Males might therefore lack the preadaptations necessary to evolve altruism in social contexts. While anecdotal observations of male contributions to colony life have been reported, there are few reports of male participation in nest defence. In apoid wasps, there have been several reports of male nest-guarding behaviour in solitary species, potentially setting the evolutionary stage for similar behaviours in social lineages. Here, we present evidence of active and effective nest defence in males of the social apoid wasp Microstigmus nigrophthalmus. Males were observed chasing intruders away from the nest, and the presence of males had a significant effect on nest survival when females were removed. Males potentially obtained direct benefits through defence, so that defence may not represent male altruism. However, our results do show that males can perform acts that benefit their colony.     

Food exploitation by a winter flock of greylag geese: behavioral dynamics,competition and social status

Summary To investigate the dynamics of the winter flock patch exploitation, feeding experiments were performed with 140 semi-tame, free-roaming greylag geese (Anser anser). Three different initial densities of barley were offered on a 50-m² patch: low (1600 grains/m²), intermediate (3300/m²) and high (I 1500/m²). Goose numbers on the patch, peck rates, and frequencies of agonistic encounters and of alert postures were observed by scanning the flock and in focal individuals. At low and intermediate initial food densities, peck rates decreased with food density, whereas at high food density, peck rates decreased only slightly over the feeding bout. Agonistic interactions increased as food decreased. A switch from exploitation to interference competition occurred at a threshold of approximately 900 remaining grains per square meter. With high initial food density, agonistic encounters began to increase gradually after 20–30 min. The same general patterns were observed for frequency of alertness. Family members occupied the patch for the longest time periods. The family female and the offspring fed most intensely. The family gander however, fed significantly less but was more aggressive and vigilant than all other social categories. Solitary geese spent their time on the patch doing little else but feeding and were the first to leave. It seems that high-ranking families and low-ranking singles adopt different competitive strategies, the former being superior interference competitors whereas the latter are forced scramble competitors.This study is dedicated to the memory of Konrad Lorenz Correspondence to: K. Kotrschal     

Exploration and exploitation of food sources by social insect colonies: a revision of the scout-recruit concept

Social insect colonies need to explore and exploit multiple food sources simultaneously and efficiently. At the individual level, this colony-level behaviour has been thought to be taken care of by two types of individual: scouts that independently search for food, and recruits that are directed by nest mates to a food source. However, recent analyses show that this strict division of labour between scouts and recruits is untenable. Therefore, a modified concept is presented here that comprises the possible behavioural states of an individual forager (novice forager, scout, recruit, employed forager, unemployed experienced forager, inspector and reactivated forager) and the transitions between them. The available empirical data are reviewed in the light of both the old and the new concept, and probabilities for the different transitions are derived for the case of the honey-bee. The modified concept distinguishes three types of foragers that may be involved in the exploration behaviour of the colony: novice bees that become scouts, unemployed experienced bees that scout, and lost recruits, i.e. bees that discover a food source other than the one to which they were directed to by their nest mates. An advantage of the modified concept is that it allows for a better comparison of studies investigating the different roles performed by social insect foragers during their individual foraging histories. Received: 29 December 1999 / Revised: 25 February 2000 / Accepted: 16 October 2000     

The nature and frequency of social interactions among free-living prairie voles (Microtus ochrogaster)

Hypotheses relating the behavior of voles to their population cycles often assume that the rate of social interaction increases with population density. To test this assumption, we examined the frequency of social interactions in a population of prairie voles (Microtus ochrogaster) over a 7-year period. In addition, we characterized space use by resident animals, patterns of visitation by nonresidents to nests, and participants in social interactions. Social groups within the population typically displayed little overlap in their use of space, even at high population densities. Nevertheless, nonresidents, particularly wandering males, were captured as visitors at nests. The number of visits per social group did not increase in a simple linear manner with population density and was particularly variable when there were fewer than 100 animals/ha. At such times, more single females and fewer pairs received visits from males than expected based on the frequency of occurrence of these groups in the population; a similar pattern was noted during periods of high population density (≥100 animals/ha) but the comparisons failed to reach statistical significance. Furthermore, at high population density, more communal groups received visits from females than expected. Patterns of visitation to communal groups were influenced by the number of adult male residents (winter only), but not by the number of adult female residents or presence of philopatric female offspring. These data indicate that the frequency of social contact in prairie voles does not increase linearly with population density and is influenced by the spacing and possible mate-guarding behavior of resident animals. Received: 7 January 1998 / Accepted after revision: 16 May 1998     

Influence of low and decreasing food levels on Daphnia-algal interactions: Numerical experiments with a new dynamic energy budget model

Based on numerical experiments with a new physiologically structured population model we demonstrate that predator physiology under low food and under starving conditions can have substantial implications for population dynamics in predator-prey interactions. We focused on Daphnia-algae interactions as model system and developed a new dynamic energy budget (DEB) model for individual daphnids. This model integrates the κ-rule approach common to net assimilation models into a net-production model, but uses a fixed allocation of net-productive energy in juveniles. The new DEB-model agrees well with the results of life history experiments with Daphnia. Compared to a pure κ-rule model the new allocation scheme leads to significant earlier maturation at low food levels and thus is in better agreement with the data. Incorporation of the new DEB-model into a physiologically structured population model using a box-car elevator technique revealed that the dynamics of Daphnia-algae interactions are highly sensitive to the assumptions on the energy allocation of juveniles under low food conditions. Additionally we show that also other energy allocation rules of our DEB-model concerning decreasing food levels and starving conditions at the individual level have strong implications for Daphnia-algae interactions at the population level. With increasing carrying capacity of algae a stable equilibrium with coexistence of Daphnia occurs and algae shifts to limit cycles. The amplitudes of the limit cycles increase with increasing percentage of sustainable weight loss. If a κ-rule energy allocation is applied to juveniles, the stable equilibrium occurs for a much narrower range of algal carrying capacities, the algal concentration at equilibrium is about 2 times larger, and the range of algae carrying capacities at which daphnids become extinct extends to higher carrying capacities than in the new DEB-model. Because predator-prey dynamics are very sensitive to predator physiology under low food and starving conditions, empirical constraints of predator physiology under these conditions are essential when comparing model results with observations in laboratory experiments or in the field.     

Almost royal: incomplete suppression of host worker ovarian development by a social parasite wasp

Insect social parasites, like other parasites, may benefit from inhibiting their host from reproducing (complete or partial parasitic castration) because they can then exploit more of the host’s resources for their own reproduction. In particular, social parasites that kill or expel the host queen need to prevent host workers from reproducing; this is a common worker response to the absence of their queen. Indeed, host workers would benefit from detecting the presence of the parasite and investing in direct and indirect fitness. Studying whether and how social parasites control host worker reproduction can provide information about the degree of integration of the parasite in the host colony and help identify factors regulating workers’ reproductive decisions in social insects. We investigated whether the paper wasp social parasite, Polistes sulcifer, suppresses Polistes dominula (host) worker reproduction as efficiently as the dominant host female does in queen-right colonies by comparing worker reproductive efforts in parasitized and non-parasitized (control) colonies. Our results show that 6 weeks after usurpation of their colony by the social parasite, parasitized workers (1) had more developed ovaries than control workers and (2) laid more eggs as soon as the opportunity arose. This reproductive readiness of parasitized workers was not apparent 2 weeks after colony usurpation. This suggests that P. dominula workers have evolved means to react to social parasitism, as occurs in some ants, and that the parasite has only limited control over host reproduction.     

Behavioural trait assortment in a social network: patterns and implications

The social fine structure of a population plays a central role in ecological and evolutionary processes. Whilst many studies have investigated how morphological traits such as size affect social structure of populations, comparatively little is known about the influence of behaviours such as boldness and shyness. Using information on social interactions in a wild population of Trinidadian guppies (Poecilia reticulata), we construct a social network. For each individual in the network, we quantify its behavioural phenotype using two measures of boldness, predator inspection tendency, a repeatable and reliably measured behaviour well studied in the context of co-operation, and shoaling tendency. We observe striking heterogeneity in contact patterns, with strong ties being positively assorted and weak ties negatively assorted by our measured behavioural traits. Moreover, shy fish had more network connections than bold fish and these were on average stronger. In other words, social fine structure is strongly influenced by behavioural trait. We assert that such structure will have implications for the outcome of selection on behavioural traits and we speculate that the observed positive assortment may act as an amplifier of selection contributing to the maintenance of co-operation during predator inspection.     

The red and the black: habituation and the dear-enemy phenomenon in two desert Pheidole ants

Many species of territorial animals are more aggressive toward strangers than neighbors, a pattern of aggression referred to as the ’dear-enemy phenomenon.’ In many cases, the mechanism by which neighbors are discriminated from strangers and the function of neighbor-stranger discrimination remain controversial. We investigated the spatial patterns of inter-colony aggression within and between two Pheidole species of seed-harvesting ants in the Mojave Desert of California by quantifying aggression between colonies in standardized staged encounters. We also tested whether the level of fighting between workers of two colonies is affected by previous exposure to each other. We show that neighbors (i.e., colonies less that 2.6 m away) of either species are treated less aggressively than more distant colonies and that habituation may be a mechanism by which this discrimination is achieved. The variation in aggression among spatially distant colonies also suggests that additional genetic or environmental factors are involved in recognition. The function of the dear-enemy phenomenon in these ant species may be related to the greater risk to the resources of a colony presented by strange workers than workers from a neighboring colony. Received: 18 November 1999 / Received in revised form: 3 April 2000 / Accepted: 3 May 2000     

The endocrinology of male rhesus macaque social and reproductive status: a test of the challenge and social stress hypotheses

Social status primarily determines male mammalian reproductive success, and hypotheses on the endocrinology of dominance have stimulated unprecedented investigation of its costs and benefits. Under the challenge hypothesis, male testosterone levels rise according to competitive need, while the social stress hypothesis predicts glucocorticoid (GC) rises in high-ranking individuals during social unrest. Periods of social instability in group-living primates, primarily in baboons, provide evidence for both hypotheses, but data on social instability in seasonally breeding species with marked social despotism but lower reproductive skew are lacking. We tested these hypotheses in seasonally breeding rhesus macaques on Cayo Santiago, Puerto Rico. We documented male fecal GC and androgen levels over a 10-month period in relation to rank, age, natal status, and group tenure length, including during a socially unstable period in which coalitions of lower ranked males attacked higher ranked males. Androgen, but not GC, levels rose during the mating season; older males had lower birth season levels but underwent a greater inter-season rise than younger males. Neither endocrine measure was related to rank except during social instability, when higher ranked individuals had higher and more variable levels of both. High-ranking male targets had the highest GC levels of all males when targeted and also had high and variable GC and androgen levels across the instability period. Our results provide evidence for both the challenge and social stress hypotheses.     

Food webs and intraguild predation: community interactions of a native mesocarnivore

Trophic level interactions between predators create complex relationships such as intraguild predation. Theoretical research has predicted two possible paths to stability in intraguild systems: intermediate predators either outcompete higher-order predators for shared resources or select habitat based on security. The effects of intraguild predation on intermediate mammalian predators such as swift foxes (Vulpes velox) are not well understood. We examined the relationships between swift foxes and both their predators and prey, as well the effect of vegetation structure on swift fox-coyote (Canis latrans) interactions, between August 2001 and August 2004. In a natural experiment created by the Pinon Canyon Maneuver Site in southeastern Colorado, USA, we documented swift fox survival and density in a variety of landscapes and compared these parameters in relation to prey availability, coyote abundance, and vegetation structure. Swift fox density varied significantly between study sites, while survival did not. Coyote abundance was positively related to the basal prey species and vegetation structure, while swift fox density was negatively related to coyote abundance, basal prey species, and vegetation structure. Our results support the prediction that, under intraguild predation in terrestrial systems, top predator distribution matches resource availability (resource match), while intermediate predator distribution inversely matches predation risk (safety match). While predation by coyotes may be the specific cause of swift fox mortality in this system, the more general mechanism appears to be exposure to predation moderated by shrub density.