SOCPROG programs: analysing animal social structures

SOCPROG is a set of programs which analyses data on animal associations. Data usually come from observations of the social behaviour of individually identifiable animals. Associations among animals, sampling periods, restrictions on the data and association indices can be defined very flexibly. SOCPROG can analyse data sets including 1,000 or more individuals. Association matrices are displayed using sociograms, principal coordinates analysis, multidimensional scaling and cluster analyses. Permutation tests, Mantel and related tests and matrix correlation methods examine hypotheses about preferred associations among individuals and classes of individual. Weighted network statistics are calculated and can be tested against null hypotheses. Temporal analyses include displays of lagged association rates (rates of reassociation following an association). Models can be fitted to lagged association rates. Multiple association measures, including measures produced by other programs such as genetic or range use data, may be analysed using Mantel tests and principal components analysis. SOCPROG also performs mark-recapture population analyses and movement analyses. SOCPROG is written in the programming language MATLAB and may be downloaded free from the World Wide Web.     

Play behavior and motor development in juvenile Belding’s ground squirrels (<Emphasis Type="Italic">Spermophilus beldingi</Emphasis>)

We evaluated the relationship between play behavior and motor development in juvenile Beldings ground squirrels (Spermophilus beldingi). We observed the social play behavior of juvenile S. beldingi during their primary play period and conducted motor skills tests at the beginning and end of the play period. We also placed different-sized boxes in juveniles natal areas during the play period and observed box climbing behavior to assess the difficulty of non-social play tasks undertaken by juveniles. Rates of social play were greater in heavy compared to light individuals, supporting the idea that juveniles must selectively allocate available energy among competing demands such as growth and play behavior. Juveniles played almost exclusively with littermates; however, they did not play at equal rates with all partners. Rather, they tended to have one preferred partner and to devote significantly less time to playing with each additional partner, indicating that stability and familiarity are important in social play interactions. Motor skill levels at the end of the play period were greater among juveniles who played with many compared to few partners, suggesting that exposure to a variety of play interactions helps facilitate motor development. Improvement in motor skills was greater among juveniles who played at high compared to low rates, had several compared to few male play partners, and engaged in several compared to few play bouts with male partners, suggesting that elements of play with male partners prominently influence motor development. The partner with whom males played at the highest rate tended to be male, and rates of social play were highest in male biased litters, raising the possibility that improvement in motor proficiency arising from play with males may be especially important for young males. The size of experimental boxes that juveniles climbed increased during the play period, supporting the idea that individuals choose play tasks that pose the greatest challenge.Communicated by T. Czeschlik     

Do great tits rely on inadvertent social information from blue tits? A habitat selection experiment

Sympatric species sharing requirements are competitors, but recent evidence suggests that heterospecifics may also be used as a source of information. The heterospecific habitat copying hypothesis proposes that individuals of one species might use information inadvertently produced by the breeding performance of individuals of other species to assess habitat quality whenever the two species share needs. In this study, we provide the first experimental test of this hypothesis by examining whether the manipulated reproductive success of blue tits (Cyanistes caeruleus) is used as heterospecific inadvertent social information (ISI) in breeding-habitat selection by sympatric great tits (Parus major). The reproductive success of blue tits was manipulated 1year at the scale of patches by transferring nestlings from decreased to increased patches. No evidence was found of great tits using the reproductive success of blue tits as a source of heterospecific ISI. However, dispersal decisions by adult great tits correlated with information on con- and heterospecific densities, which constitute other sources of ISI. As density and breeding performance are tightly intertwined forms of information, the difficulty in distinguishing between them might lead great tits to use heterospecific ISI more in the form of density than breeding performance when making dispersal decisions.     

Dual mechanism of queen influence over sex ratio in the ant Pheidole pallidula

Social Hymenoptera are general models for the study of parent-offspring conflict over sex ratio, because queens and workers frequently have different reproductive optima. The ant Pheidole pallidula shows a split distribution of sex ratios with most of the colonies producing reproductives of a single sex. Sex ratio specialization is tightly associated with the breeding system, with single-queen (monogynous) colonies producing male-biased brood and multiple-queen (polygynous) colonies female-biased brood. Here, we show that this sex specialization is primarily determined by the queens influence over colony sex ratio. Queens from monogynous colonies produce a significantly more male-biased primary sex ratio than queens from polygynous colonies. Moreover, queens from monogynous colonies produce a significantly lower proportion of diploid eggs that develop into queens and this is associated with lower rate of juvenile hormone (JH) production compared to queens from polygynous colonies. These results indicate that queens regulate colony sex ratio in two complementary ways: by determining the proportion of female eggs laid and by hormonally biasing the development of female eggs into either a worker or reproductive form. This is the first time that such a dual system of queen influence over colony sex ratio is identified in an ant.     

Association networks in spider monkeys (<Emphasis Type="Italic">Ateles geoffroyi</Emphasis>)

We use two novel techniques to analyze association patterns in a group of wild spider monkeys (Ateles geoffroyi) studied continuously for 8 years. Permutation tests identified association rates higher or lower than chance expectation, indicating active processes of companionship and avoidance as opposed to passive aggregation. Network graphs represented individual adults as nodes and their association rates as weighted edges. Strength and eigenvector centrality (a measure of how strongly linked an individual is to other strongly linked individuals) were used to quantify the particular role of individuals in determining the network's structure. Female–female dyads showed higher association rates than any other type of dyad, but permutation tests revealed that these associations cannot be distinguished from random aggregation. Females formed tightly linked clusters that were stable over time, with the exception of immigrant females who showed little association with any adult in the group. Eigenvector centrality was higher for females than for males. Adult males were associated mostly among them, and although their strength of association with others was lower than that of females, their association rates revealed a process of active companionship. Female–male bonds were weaker than those between same-sex pairs, with the exception of those involving young male adults, who by virtue of their strong connections both with female and male adults, appear as temporary brokers between the female and male clusters of the network. This analytical framework can serve to develop a more complete explanation of social structure in species with high levels of fission–fusion dynamics. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James)     

Insights on common dolphin (<Emphasis Type="Italic">Delphinus delphis</Emphasis>) social organization from genetic analysis of a mass-stranded pod

Compared to terrestrial mammals, little is known of cetacean social systems as they are generally less accessible to behavioral investigations due to their aquatic environment. The present study investigates group structure of the pelagic common dolphin, Delphinus delphis, using genetic markers. Tissue samples from 52 individuals representing a recent live mass-stranding event were compared to 42 single strandings taken from presumably different groups. The mass-stranding event occurred in 2002 on the French coast of the English Channel, whereas the single strandings were collected between 1993 and 2003 along the western coast of France (Bay of Biscay and English Channel). Analysis of mitochondrial DNA control region sequences indicated that genetic variability within the mass-stranded pod was similar to variability observed in single strandings. The mass-stranded group was composed of 41 different mitochondrial haplotypes or matrilines while the single strandings revealed 29 different haplotypes. Analysis of 11 microsatellite loci revealed that average relatedness of the mass-stranded pod was not different from average relatedness among all single strandings suggesting that individuals within the group had no closer kin relationships than animals taken from presumably different groups. These results do not support a matriarchal system and suggest that common dolphins constituting a pod are not necessarily genetically related.     

Reputation is valuable within and outside one’s own social group

To find conditions under which humans cooperate within groups of unrelated individuals has been of major interest in the behavioral sciences. The experimental paradigm for studying potential cooperation in social dilemmas is the public goods game. Here humans regularly fail to sustain a public resource cooperatively. However, the need to maintain good reputation for other social interactions, such as indirect reciprocity, has been identified as an effective mechanism to sustain cooperation in public goods situations. As a side effect of building a good reputation through cooperative actions, an individual provides direct benefits to members of his/her own social group. These benefits could be an incentive to reward a good reputation of group members. Here we show experimentally that building a good reputation through cooperative behavior in a public goods situation is rewarded in future social interactions, not only within ones own social group but also, at a similar level, in other social groups: humans regard cooperative behavior of others as an honest signal irrespective of past direct personal benefits. Reputation gained within as well as outside ones own social group can be a driving force for selfish individuals to cooperate in public goods situations, and thereby sustain any public resource.Communicated by T. Czeschlik