Allocare in a nocturnal primate: data on the spectral tarsier, Tarsius spectrum

Non-maternal infant care in many of the small-bodied New World primate species has been hypothesized by some researchers to be related to the high infant/adult weight ratio found in these species. The spectral tarsier, Tarsius spectrum, an Old World primate, has one of the highest infant/adult weight ratios of any primate, with infants weighing between 20-33% of adult weight at birth. On the basis of the hypothesized relationship between allocare and the infant/adult weight ratio, it is predicted that the spectral tarsier will also exhibit extensive allocaretaking behaviour. The results of this study indicate that although spectral tarsiers show care by male and female subadults as well as adult males, it is extremely limited compared to the extensive allocaretaking behavior observed in New World primate species such as Aotus and Callicebus. Spectral tarsier subadult females provide substantially more allocare to infants than do subadult males or adult males. Female subadults were observed sharing food, transporting, grooming, playing, alarm calling, baby-sitting and maintaining physical contact with infants more than other age/sex classes. Although the amount of allocare exhibited by adult males and subadult males was much less than that exhibited by female subadults, the data suggest that adult and subadult male spectral tarsiers do play a small part in the care and socialization of the infant. Adult males and subadult males were both observed occasionally engaging in allocaretaking behaviors such as grooming and playing, as well as frequently patrolling and defending the territory's boundaries. The results from this study suggest that although a high infant/adult weight ratio may be a prerequisite for selection to favor extensive allocare, it is not a causal factor. Additional research is needed in order to understand better the selective pressures involved and the costs and benefits of providing allocare to subadults and adult male spectral tarsiers.     

Calves as social hubs: dynamics of the social network within sperm whale units

It is hypothesized that the primary function of permanent social relationships among female sperm whales (Physeter macrocephalus) is to provide allomothers for calves at the surface while mothers make foraging dives. In order to investigate how reciprocity of allocare within units of sperm whales facilitates group living, we constructed weighted social networks based on yearly matrices of associations (2005–2010) and correlated them across years, through changes in age and social role, to study changes in social relationships within seven sperm whale units. Pairs of association matrices from sequential years showed a greater positive correlation than expected by chance, but as the time lag increased, the correlation coefficients decreased. Over all units considered, calves had high values for all measured network statistics, while mothers had intermediate values for most of the measures, but high values for connectedness and affinity. Mothers showed sharp drops in strength and connectedness in the first year of their new calves'' lives. These broad patterns appear to be consistent across units. Calves appeared to be significant nodes in the network of the social unit, and thus provide quantitative support for the theory in which communal care acts as the evolutionary force behind group formation in this species.     

The effects of residential locality on parental and alloparental investment among the Aka foragers of the central African Republic

In this paper I examine the intracultural variability of parental and alloparental caregiving among the Aka foragers of the Central African Republic. It has been suggested that maternal kin offer higher frequencies of allocare than paternal kin and that maternal investment in infants will decrease when alloparental assistance is provided. Behavioral observations were conducted on 15 eight- to twelve-monthold infants. The practice of brideservice and the flexibility of Aka residence patterns offered a means to test the effect of maternal residence on parental and alloparental investment. There was significant variation in the frequency of investment and who supplied care to infants depending on whether mothers resided with their kin or their husbands’ kin. However, in spite of the variation in allocare, when all categories of caregivers were examined collectively, infants received similar overall levels of care. Courtney L. Meehan is a Ph.D. student in cultural anthropology at Washington State University. Her research interests include parenting, alloparenting, female social networks, and female-female cooperation and competition.     

The evolution of non-maternal care in anthropoid primates: a test of the hypotheses

The amount of non-maternal care (allocare) found in primates varies greatly from species to species. Our paper examines this variation and focuses on possible reasons why mothers in some anthropoid primate species are prepared to relinquish their infants to other carers whereas others are not. We use data collected from the literature to test a number of hypotheses that attempt to explain the observed variation in non-maternal care. Analyses were carried out using comparative methods that control for the influence of both body size and phylogeny on life-history and ecological variation. The results support previous studies of both primates and other mammals in finding a clear link between the amount of allocare provided and female reproductive rates. Species with high allocare levels grow rapidly post-natally and wean their infants at a younger age (but at the same relative size) than species of the same body size with lower allocare levels. This early weaning allows high allocare species to support higher birth rates than low allocare species but does not result in earlier maturation. Our results, therefore, suggest that mothers allow non-maternal care of their infants in order to increase their own reproductive output. It is not clear whether such a strategy also benefits the infant, as we could find no link between the presence of allocare and early maturation (which would increase the infant's reproductive output) or between allocare levels and infant survival (as measured by vulnerability to infanticide). This suggests that mothers and infants might come into conflict over transfer to other carers, as the benefits to the mother may not be accompanied by benefits to the infant. However, although mothers may benefit from allocare in some circumstances, they will not be expected to allow allocare if the costs are high (e.g. if there is a high risk to the infant).     

西藏小昌都黑白仰鼻猴的雄婴照料:能量胁迫导致?

雄猴照料婴猴行为通常都是从雄猴的角度来分析该行为具有何种适应功能,很少有研究关心为什么雌猴会让雄猴参与照料活动。该文通过西藏小昌都黑白仰鼻猴群的系统观察取样,采用回归分析雄婴照料行为季节变动与婴猴年龄及生态因子（环境温度、食物供应）之间的关系,试图分析是否由于能量胁迫导致雌猴让雄猴参与照料活动,并通过分析高能量胁迫季节（环境温度低、食物缺乏,婴猴年幼需要雌猴携带多）与低能量胁迫季节（环境温度高、食物丰富,婴猴年长需要雌猴携带少）雄婴照料行为差异检验了能量胁迫假说。结果表明雄婴照料行为与温度呈负相关函数,而雌猴照料婴猴行为与温度呈正相关函数;在高能量胁迫的季节,雄猴参与照料行为多,而在低能量胁迫的季节雄猴照料行为少。因此,作者认为是能量胁迫迫使雌猴放弃部分照料婴猴的机会,雄猴利用照料机会获得了其他利益。     

Who Responds to Crying?

!Kung San (Bushman) hunter-gatherers have unusually high levels of mother-infant contact and represent one of the environments of human evolutionary adaptedness (EEAs). Studies among the !Kung show that levels of crying—the most basic sign of mammalian infant distress—are low, and response to crying is high, and some suggest that responses are overwhelmingly maternal. We show that although !Kung mothers respond to crying most often, one-third of crying bouts are managed solely by someone else. Mothers responded to all bouts lasting ≥30 s, but in half of these responses they were joined by one or more others. Mothers are the most consistent responders, but multiple caregiving is common. The mother is rarely alone when her baby cries; others often substitute or join her in interventions. This social support may facilitate the high levels of maternal responsiveness characteristic of the !Kung, and of hunter-gatherers generally, but it is also consistent with recent theory emphasizing nonmaternal care (allocare) and cooperative breeding.     

Early Social Networks Predict Survival in Wild Bottlenose Dolphins

A fundamental question concerning group-living species is what factors influence the evolution of sociality. Although several studies link adult social bonds to fitness, social patterns and relationships are often formed early in life and are also likely to have fitness consequences, particularly in species with lengthy developmental periods, extensive social learning, and early social bond-formation. In a longitudinal study of bottlenose dolphins (Tursiops sp.), calf social network structure, specifically the metric eigenvector centrality, predicted juvenile survival in males. Additionally, male calves that died post-weaning had stronger ties to juvenile males than surviving male calves, suggesting that juvenile males impose fitness costs on their younger counterparts. Our study indicates that selection is acting on social traits early in life and highlights the need to examine the costs and benefits of social bonds during formative life history stages.     

Inferring a District-Based Hierarchical Structure of Social Contacts from Census Data

Researchers have recently paid attention to social contact patterns among individuals due to their useful applications in such areas as epidemic evaluation and control, public health decisions, chronic disease research and social network research. Although some studies have estimated social contact patterns from social networks and surveys, few have considered how to infer the hierarchical structure of social contacts directly from census data. In this paper, we focus on inferring an individual’s social contact patterns from detailed census data, and generate various types of social contact patterns such as hierarchical-district-structure-based, cross-district and age-district-based patterns. We evaluate newly generated contact patterns derived from detailed 2011 Hong Kong census data by incorporating them into a model and simulation of the 2009 Hong Kong H1N1 epidemic. We then compare the newly generated social contact patterns with the mixing patterns that are often used in the literature, and draw the following conclusions. First, the generation of social contact patterns based on a hierarchical district structure allows for simulations at different district levels. Second, the newly generated social contact patterns reflect individuals social contacts. Third, the newly generated social contact patterns improve the accuracy of the SEIR-based epidemic model.     

Variation in the Social Systems of Extant Hominoids: Comparative Insight into the Social Behavior of Early Hominins

The observed social systems of extant apes and humans suggest that the common ancestral state for Miocene hominoids was living in multimale–multifemale groups that exhibited a tendency to fission and fusion in response to ecological and/or social variables. The Hominoidea share a set of social commonalities, notably a social niche that extends beyond kin and beyond the immediate social group, as well as extensive intraspecific flexibility in social organization. We propose that an essential feature of hominoid evolution is the shift from limited plasticity in a generalized social ape to expanded behavioral plasticity as an adaptive niche. Whereas in most nonhominoid primates variability and flexibility take the shape of specific patterns of demographic flux and interindividual relationships, we can consider behavioral flexibility and plasticity as a means to an end in hominoid socioecological landscapes. In addition, the potential for innovation, spread, and inheritance of behavioral patterns and social traditions is much higher in the hominoids, especially the great apes, than in other anthropoid primates. We further suggest that this pattern forms a basis for the substantial expansion of social complexity and adaptive behavioral plasticity in the hominins, especially the genus Homo. Our objectives in this article are threefold: 1) summarize the variation in the social systems of extant hominoid taxa; 2) consider the evolutionary processes underlying these variations; and 3) expand upon the traditional socioecological model, especially with respect to reconstructions of early hominin social behavior. We emphasize a central role for both ecological and social niche construction, as well as behavioral plasticity, as basal hominoid characteristics. Over evolutionary time these characteristics influence the patterns of selection pressures and the resulting social structures. We propose that a mosaic of ecological and social inheritance patterns should be considered in the reconstruction of early hominin social systems.     

Effect of rearing style on the development of social behaviour in young ravens (Corvus corax)

Early social experiences can affect the development and expression of individual social behaviour throughout life. In particular, early-life social deprivations, notably of parental care, can later have deleterious consequences. We can, therefore, expect rearing procedures such as hand-raising—widely used in ethology and socio-cognitive science—to alter the development of individual social behaviour. We investigated how the rearing style later affected (a) variation in relationship strength among peers and (b) individuals’ patterns of social interactions, in three captive groups of juvenile non-breeders consisting of either parent-raised or hand-raised birds, or a mix of both rearing styles. In the three groups, irrespectively of rearing style: strongest relationships (i.e., higher rates of association and affiliations) primarily emerged among siblings and familiar partners (i.e., non-relatives encountered in early life), and mixed-sex and male–male partners established relationships of similar strength, indicating that the rearing style does not severely affect the quality and structure of relationships in young ravens. However, compared to parent-raised ravens, hand-raised ravens showed higher connectedness, i.e., number of partners with whom they mainly associated and affiliated, but formed on average relationships of lower strength, indicating that social experience in early life is not without consequences on the development of ravens’ patterns of social interaction. The deprivation of parental care associated with the presence of same-age peers during hand-raising seemed to maximize ravens’ propensity to interact with others, indicating that besides parents, interactions with same-age peers matter. Opportunities to interact with, and socially learn from peers, might thus be the key to the acquisition of early social competences in ravens.     

Prolactin levels of fathers and helpers related to alloparental care in common marmosets, Callithrix jacchus

Previous studies have suggested that prolactin may play a role in regulating allocare behaviour in cotton-top tamarins, Saguinus oedipus. In this study, we investigate the prolactin profile of 3 groups of captive common marmosets, Callithrix jacchus. Carrying behaviour in this species was observed after parturition. Prolactin assays of blood samples of both fathers and helpers (sub-adult non-fathers) in 3 family groups were taken for 8 weeks before and after birth of the infants. The after-birth condition was divided into 2 groups: carrying and non-carrying animals. The results suggest a relationship between prolactin levels and allocare behaviour, with carrying behaviour being associated with increased prolactin in both fathers and helpers. This suggests that extra prolactin is produced in response to physical contact, and may be associated with carrying behaviour. Also, prolactin production may be related to learning parental skills in Neotropical primates.     

Social organization, reproduction and rearing strategies of Callimico goeldii: new clues from the wild

The callitrichines are a specialized radiation of primates that are characterized in part by variable social systems and cooperative infant care. Callimico goeldii, unlike the other callitrichines, have single rather than twin offspring, reducing the need for allocare and permitting synchronous breeding within groups. Low mortality rates among offspring and unstable social groups are suggested to be possible factors that have led to single births among C. goeldii. Single offspring may benefit from greater maternal investment and more frequent food sharing than twin offspring, factors that may help to explain why C. goeldii reaches sexual maturity more rapidly than other callitrichines. In addition, increased breeding opportunities for young C. goeldii females may have selected for rapid maturation rates among this species. Postpartum ovulation and aseasonal resource availability appear to permit females to have biannual birth seasons, further increasing the potential reproductive output.     

Inferring the Structure of Social Contacts from Demographic Data in the Analysis of Infectious Diseases Spread

Social contact patterns among individuals encode the transmission route of infectious diseases and are a key ingredient in the realistic characterization and modeling of epidemics. Unfortunately, the gathering of high quality experimental data on contact patterns in human populations is a very difficult task even at the coarse level of mixing patterns among age groups. Here we propose an alternative route to the estimation of mixing patterns that relies on the construction of virtual populations parametrized with highly detailed census and demographic data. We present the modeling of the population of 26 European countries and the generation of the corresponding synthetic contact matrices among the population age groups. The method is validated by a detailed comparison with the matrices obtained in six European countries by the most extensive survey study on mixing patterns. The methodology presented here allows a large scale comparison of mixing patterns in Europe, highlighting general common features as well as country-specific differences. We find clear relations between epidemiologically relevant quantities (reproduction number and attack rate) and socio-demographic characteristics of the populations, such as the average age of the population and the duration of primary school cycle. This study provides a numerical approach for the generation of human mixing patterns that can be used to improve the accuracy of mathematical models in the absence of specific experimental data.     

Circulation of type 1 vaccine-derived poliovirus in the Philippines in 2001

In 2001, highly evolved type 1 circulating vaccine-derived poliovirus (cVDPV) was isolated from three acute flaccid paralysis patients and one contact from three separate communities in the Philippines. Complete genomic sequencing of these four cVDPV isolates revealed that the capsid region was derived from the Sabin 1 vaccine strain but most of the noncapsid region was derived from an unidentified enterovirus unrelated to the oral poliovirus vaccine (OPV) strains. The sequences of the cVDPV isolates were closely related to each other, and the isolates had a common recombination site. Most of the genetic and biological properties of the cVDPV isolates were indistinguishable from those of wild polioviruses. However, the most recently identified cVDPV isolate from a healthy contact retained the temperature sensitivity and partial attenuation phenotypes. The sequence relationships among the isolates and Sabin 1 suggested that cVDPV originated from an OPV dose given in 1998 to 1999 and that cVDPV circulated along a narrow chain of transmission. Type 1 cVDPV was last detected in the Philippines in September 2001, and population immunity to polio was raised by extensive OPV campaigns in late 2001 and early 2002.     

Dynamic contact stress patterns on the tibial plateaus during simulated gait: A novel application of normalized cross correlation

The spatial distribution and pattern of local contact stresses within the knee joint during activities of daily living have not been fully investigated. The objective of this study was to determine if common contact stress patterns exist on the tibial plateaus of human knees during simulated gait. To test this hypothesis, we developed a novel normalized cross-correlation (NCC) algorithm and applied it to the contact stresses on the tibial plateaus of 12 human cadaveric knees subjected to multi-directional loads mimicking gait. The contact stress profiles at different locations on the tibial plateaus were compared, where regions with similar contact stress patterns were identified across specimens. Three consistent regional patterns were found, among them two most prominent contact stress patterns were shared by 9–12 of all the knees and the third pattern was shared by 6–8 knees. The first pattern was located at the posterior aspect of the medial tibial plateau and had a single peak stress that occurred during the early stance phase. The second pattern was located at the central-posterior aspects of the lateral plateau and consisted of two peak stresses coincident with the timing of peak axial force at early and late stance. The third pattern was found on the anterior aspect of cartilage-to-cartilage contact region on the medial plateau consisted of double peak stresses. The differences in the location and profile of the contact stress patterns suggest that the medial and lateral menisci function to carry load at different points in the gait cycle: with the posterior aspect of the medial meniscus consistently distributing load only during the early phase of stance, and the posterior aspect of the lateral meniscus consistently distributing load during both the early and late phases of stance. This novel approach can help identify abnormalities in knee contact mechanics and provide a better understanding of the mechanical pathways leading to post-traumatic osteoarthritis.     

Social Play in the Domestic Cat

The social play of domestic cats in the laboratory and at homewas examined. Categories and sequences of motor patterns wereidentified and analyzed. The developmental period during whichsocial play was most frequent was established to be from 4 weeksto 4 months. The results suggest that social play functionsto provide specific forms of exercise and as a means of developingand maintaining social relations among littermates. The declinein social play appears to be related to dispersal of the youngand to a decrease in preference for social contact.     

Society, demography and genetic structure in the spotted hyena

Spotted hyenas (Crocuta crocuta) are large mammalian carnivores, but their societies, called 'clans', resemble those of such cercopithecine primates as baboons and macaques with respect to their size, hierarchical structure, and frequency of social interaction among both kin and unrelated group-mates. However, in contrast to cercopithecine primates, spotted hyenas regularly hunt antelope and compete with group-mates for access to kills, which are extremely rich food sources, but also rare and ephemeral. This unique occurrence of baboon-like sociality among top-level predators has favoured the evolution of many unusual traits in this species. We briefly review the relevant socio-ecology of spotted hyenas, document great demographic variation but little variation in social structure across the species' range, and describe the long-term fitness consequences of rank-related variation in resource access among clan-mates. We then summarize patterns of genetic relatedness within and between clans, including some from a population that had recently gone through a population bottleneck, and consider the roles of sexually dimorphic dispersal and female mate choice in the generation of these patterns. Finally, we apply social network theory under varying regimes of resource availability to analyse the effects of kinship on the stability of social relationships among members of one large hyena clan in Kenya. Although social bonds among both kin and non-kin are weakest when resource competition is most intense, hyenas sustain strong social relationships with kin year-round, despite constraints imposed by resource limitation. Our analyses suggest that selection might act on both individuals and matrilineal kin groups within clans containing multiple matrilines.     

Lack of congruence of genetic and niche divergence in Podarcis hispanicus complex

Niche divergence among closely related lineages can be informative on the ecological and evolutionary processes involved in differentiation, particularly in the case of cryptic species complexes. Here we compared phylogenetic relationships and niche similarity between pairs of lineages included in the Podarcis hispanicus complex to examine patterns of niche divergence and its role in the organization of current diversity patterns, as allopatric, parapatric, and sympatric lineages occur in the Western Mediterranean Basin. First, we used ecological niche models to characterize the realized climatic niche of each Podarcis hispanicus complex lineage based on topographic and climatic variables, to identify important variables, and to test for niche conservatism or divergence between pairs of lineages. Variables related to precipitation generally exhibited the highest contribution to niche models, highlighting the importance of rainfall levels in shaping distributions of Podarcis wall lizards. We found that most forms have significant differences in realized climatic niches that do not follow the pattern of mitochondrial divergence. These results lend support to the hypothesis that genetic divergence across Podarcis hispanicus complex most likely occurred in allopatric conditions, mostly with significant niche divergence. Competition after secondary contact is also suggested by the common occurrence of niche overlap between lineages that exhibit strictly parapatric distribution. The almost continuous distribution of Podarcis lizards in the study area appears to be a result of a combination of complementary suitable niches and competition, which seem two important mechanisms limiting geographic distributions and restricting the existence of extensive contact zones.     

Intragroup Social Relationships of Male Alouatta palliata on Barro Colorado Island,Republic of Panama

We examined social and spatial relations of adult males in one group of mantled howling monkeys (Alouatta palliata) on Barro Colorado Island (BCI) in central Panama to document patterns of association. Beyond the existence of an alpha male, we could not distinguish any linear dominance hierarchy among the 6 study males. All males copulated with estrous females. Our findings contrast with reports of intragroup male behavior in Costa Rican howlers. Study males engaged in little or no affiliative or agonistic behavior with one another, but engaged in significantly more such interactions with females. The alpha male, the oldest male and a younger male were most frequently in association with females. Of group males, the oldest male associated significantly more with other males. Overall, male behavior in mantled howling monkeys on BCI generally followed the van Hooff and van Schaik (1994) model of male relationships. The low incidence of intragroup social interactions of any type in the focal males may reflect the energetic costs of social behavior. We suggest that intragroup social relationships among mantled howler males are structured by more subtle means than overt physical interactions, possibly including vocal communication, relationships with individual group females, and kinship.     

Dominance and the social behavior of adult female bonnet macaques (Macaca radiata)

Female bonnet macaques are ranked according to naturally occurring dyadic aggressive interactions. Several possible correlates of these ranks are examined. Maternal rank and early life history are found to be highly correlated with rank. In addition, a number of social relationships (female/female presentations, female/female mounts, contact among females and aggression) are found to be related to rank. The meaning of rank outside of head to head encounters is discussed. This research was supported in part by the California Primate Research Center, University of California, Davis, Davis, California 95616.