Relatedness facilitates cooperation in the subsocial spider, Stegodyphus tentoriicola

Background  

Cooperative hunting and foraging in spiders is rare and prone to cheating such that the actions of selfish individuals negatively affect the whole group. The resulting social dilemma may be mitigated by kin selection since related individuals lose indirect fitness benefits by acting selfishly. Indeed, cooperation with genetic kin reduces the disadvantages of within-group competition in the subsocial spider Stegodyphus lineatus, supporting the hypothesis that high relatedness is an important pre-adaptation in the transition to sociality in spiders. In this study we examined the consequences of group size and relatedness on cooperative feeding in the subsocial spider S. tentoriicola, a species suggested to be at the transition to permanent sociality.     

A Quantitative Index of Sociality and Its Application to Group‐Living Spiders and Other Social Organisms

Species are often classified in discrete categories, such as solitary, subsocial, social and eusocial based on broad qualitative features of their social systems. Often, however, species fall between categories or species within a category may differ from one another in ways that beg for a quantitative measure of their sociality level. Here, we propose such a quantitative measure in the form of an index that is based on three fundamental features of a social system: (1) the fraction of the life cycle that individuals remain in their social group, (2) the proportion of nests in a population that contain multiple vs. solitary individuals and (3) the proportion of adult members of a group that do not reproduce, but contribute to communal activities. These are measures that should be quantifiable in most social systems, with the first two reflecting the tendencies of individuals to live in groups as a result of philopatry, grouping tendencies and intraspecific tolerance, and the third potentially reflecting the tendencies of individuals to exhibit reproductive altruism. We argue that this index can serve not only as a way of ranking species along a sociality scale, but also as a means of determining how level of sociality correlates with other aspects of the biology of a group of organisms. We illustrate the calculation of this index for the cooperative social spiders and the African mole‐rats and use it to analyse how sex ratios and interfemale spacing correlate with level of sociality in spider species in the genus Anelosimus.     

Within‐population variation in social strategies characterize the social and mating system of an Australian lizard,Egernia whitii

The lizard genus Egernia has been suggested as an excellent model system for examining the evolution of sociality as it exhibits considerable diversity in social organization both between and within species. To date the majority of work examining the factors responsible for the evolution of sociality within Egernia has advocated a broad scale approach; identifying the social structure of specific species or populations and comparing the degree of sociality between them. However, we argue that significant advancements could also be gained by examining variation in social strategies within populations. Here we integrate a detailed, 3‐year, field‐based examination of social spacing and juvenile dispersal with molecular analyses of paternity to determine the social and mating system of a Tasmanian population of White's skink (Egernia whitii). We show that E. whitii live in small stable family groups consisting of an adult male, his female partner(s), as well as juvenile or sub‐adults individuals. In addition, while the mating system is characterized by considerable genetic monogamy, extra‐pair fertilizations are relatively common, with 34% of litters containing offspring sired by males from outside the social group. We also show that traits related to social organization (social group composition, group size, stability and the level of extra‐pair paternity) vary both between and within individuals. We suggest that ecological factors, such as habitat saturation, quality and availability, play a key role in maintaining between individual variation in social strategies, and that examining these individual level processes will allow us to more clearly understand variation in sociality among species.     

The evolution of social inbreeding mating systems in spiders: limited male mating dispersal and lack of pre‐copulatory inbreeding avoidance in a subsocial predecessor

Cooperation and group living are extremely rare in spiders and only few species are known to be permanently social. Inbreeding is a key characteristic of social spiders, resulting in high degrees of within‐colony relatedness that may foster kin‐selected benefits of cooperation. Accordingly, philopatry and regular inbreeding are suggested to play a major role in the repeated independent origins of sociality in spiders. We conducted field observations and laboratory experiments to investigate the mating system of the subsocial spider Stegodyphus tentoriicola. The species is suggested to resemble the ‘missing link’ in the transition from subsociality to permanent sociality in Stegodyphus spiders because its social period is prolonged in comparison to other subsocial species. Individuals in our two study populations were spatially clustered around maternal nests, indicating that clusters consist of family groups as found in the subsocial congener Stegodyphus lineatus. Male mating dispersal was limited and we found no obvious pre‐copulatory inbreeding avoidance, suggesting a high likelihood of mating with close kin. Rates of polygamy were low, a pattern ensuring high relatedness within broods. In combination with ecological constraints, such as high costs of dispersal, our findings are consistent with the hypothesis that the extended social period in S. tentoriicola is accompanied with adaptations that facilitate the transition towards permanent sociality. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 851–859.     

Variability in the estimation of ungulate group sizes complicates ecological inference

1. Foundational work has examined adaptive social behavior in animals in relation to the costs and benefits of group living. Within this context, a “group” of animals represents an organizational unit that is integral to the study of animal ecology and evolution.
2. Definitions of animal group sizes are often subjective with considerable variability within and across species. However, investigations of both the extent and implications of such variability in the estimation of animal group sizes are currently lacking.
3. Selecting ungulates as a case study, we conducted a literature review to assess prevailing practices used to determine group sizes among terrestrial Cetartiodactyla and Perissodactyla. Via this process, we examined group size definitions for 61 species across 171 peer‐reviewed studies published between 1962 and 2018.
4. These studies quantified group sizes via estimation of ungulate aggregations in space and time. Spatial estimates included a nearest neighbor distance ranging from 1.4 m to 1,000 m, and this variation was partially explained by a weak positive correlation (|r| = .4, p < .003) with the body size of the ungulate research subjects. The temporal extent over which group size was estimated was even broader, ranging from three minutes to 24 hr.
5. The considerable variability in ungulate group size estimation that we observed complicates efforts to not only compare and replicate studies but also to evaluate underlying theories of group living. We recommend that researchers: (a) clearly describe the spatiotemporal extents over which they define ungulate group sizes, (b) highlight foundational empirical and ecological rationale for these extents, and (c) seek to align such extents among individual species so as to facilitate cross‐system comparisons of ungulate group size dynamics. We believe an integrative approach to ungulate group size estimation would readily facilitate replication, comparability, and evaluation of competing hypotheses examining the tradeoffs of animal sociality.

     

Correlates of reproductive success among field colonies of Bombus lucorum: the importance of growth and parasites

Abstract.

• 1 In natural populations, colonies of bumble bees vary in many important life history traits, such as colony size and age at maturity, or the number and sex of reproductives produced. We investigated how the presence of parasites in field populations of the bumble bee Bombus lucorum L. relates to variation in life history traits and reproductive performance. A total of thirty-six colonies was placed in accessible nest sites in the field and monitored at regular intervals throughout a season.
• 2 Among the life history correlates, early nest foundation was strongly associated with large maximum colony size, old age and large size at maturity, and this in turn with successful production of males and queens, as well as with the number of sexuals produced. Overall, reproductive success was highly skewed with only five colonies producing all the queens. Sixteen colonies failed to reproduce altogether.
• 3 The social parasite Psithyrus was abundant early in the Bombus colony cycle and preferentially invaded host nests with many first brood workers and thus disproportionately large size, i.e. those colonies that would otherwise be more likely to reproduce or produce (daughter) queens rather than males. To prevent nest loss, Psithyrus had to be removed soon after invasion. Therefore, the effects reported here can only be crude estimates.
• 4 Parasitoid conopid flies are likely to cause heavy worker mortality when sexuals are reared by the colonies. Their inferred effect was a reduction in biomass that could be invested in sexuals as well as a shift in the sex ratio at the population level resulting from failure to produce queens. We suggest to group the inferred correlates into ‘early events’ surrounding colony initiation and social parasitism, and ‘late events’ surrounding attained colony size in summer and parasitism by conopid flies. Our evidence thus provides a heuristic approach to understand the factors that affect reproductive success of Bombus colonies.

     

Environmental filtering of foraging strategies mediates patterns of coexistence in the fire ants Solenopsis geminata and Solenopsis xyloni,and their interspecific hybrids

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Female transfer in primates

Intergroup transfer by males is nearly universal among social primates. Furthermore, among the most frequently studied monkeys-savanna baboons and Japanese and rhesus macaques—females typically remain in their natal groups, so troops are composed of related matrilines. These facts strongly support two major theories: (l) that kin selection is a powerful force in patterning sociality (if one is to live in a group, one should prefer a group of one’s relatives); and (2) that the ultimate explanation for intergroup transfer is the avoidance of inbreeding depression (though both sexes would prefer to live with kin, one sex has to disperse to avoid inbreeding and for a variety of reasons the losing sex is generally male). Substantial rates of transfer by females in social species with routine male transfer would cast doubt on both ideas. In fact, evidence reviewed here indicates that female transfer is not unusual and among folivorous primates (e.g., Alouatta,the Colobinae) it seems to be routine. In addition to casting doubt on the demographic significance of inbreeding avoidance and favoring mutualistic and/or game theory interpretations of behavior over nepotistic ones, this finding supports the hypothesis that predator detection is the primary selective pressure favoring sociality for many primates. Finally, while female bonding [sensuWrangham, R. W. (1980), Behaviour75:262–299] among primates appears to be less common than generally believed, the observed correlation between female transfer and morphological adaptations to folivory provides empirical support for Wrangham’s model for the evolution of female-bonded groups.     

Nesting biology,life cycle,and interactions between females of Manuelia postica,a solitary species of the Xylocopinae (Hymenoptera: Apidae)

Abstract

The Xylocopinae contains four tribes with species which show a range of nesting habits, from solitary to social. The Manueliini is the sister group to all other Xylocopine tribes, with one genus, Manuelia, of three species found mainly in Chile. This is a solitary genus, whose biology is scarcely known for two species, M. gayatina and M. gayi, and so far completely unknown for M. postica. This paper reports on nesting substrates, nest architecture, nesting behaviours, life cycle, and interactions between females at nesting sites, for M. postica. The results indicate that M. postica presents some features which are typical of solitary life, and also some features which are unusual in solitary bees but have been reported in phylogenetically more apical social species. Our findings open interesting questions on the ecological scenarios involved in the evolution of sociality within the Xylocopinae.     

The slow—fast life style characteristics in a suite of six species of odonate larvae

• 1 The validity of the slow—fast lifestyle dichotomy proposed by Sih (1987) was tested in a suite of six odonate species from a restricted geographical area. Data on activity and microhabitat use were obtained in a laboratory study. Further necessary information on life history, macrohabitat (ephemeral‐permanent) use and vulnerability to fish predation was provided by a literature survey.
• 2 Activity was estimated as number of moves and distance moved for the six odonate larvae. Aeschna juncea, Lestes sponsa and Sympetrum danae were categorised as high‐active species, whereas Coenagrion hastulatum, Cordulia aenea and Leucorhinia dubia were categorised as low‐active species.
• 3 C. hastulatum and L. sponsa exploited microhabitats close to the water surface, C. aenea and L. dubia close to the bottom, and A. juncea and S. danae were intermediate in their water depth utilisation.
• 4 A principal component analysis of the data from the laboratory experiment and the literature survey supported the slow—fast life style dichotomy since the variables activity, macrohabitat use, life cycle length and sensitivity to fish predators were highly correlated.

     

Assessing trap and lure effectiveness for the monitoring of Sirex noctilio

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A review of the social behaviour of the giraffe Giraffa camelopardalis: a misunderstood but socially complex species

1. Until around 2000, giraffe Giraffa camelopardalis were believed to have no social structure. Despite a resurgence of interest in giraffe behaviour in around 2010, most studies are of isolated populations, making it difficult to draw general conclusions. Although it is now well established that giraffe social organisation is non-random, there is little consensus as to what influences preferred and avoided associations or the underpinning mechanisms.
2. We test two hypotheses: first, giraffe have a complex cooperative social system, exhibited by 1) stable groups of females, 2) offspring that stay in their natal group for part or all of their lives, 3) support by non-mothers in rearing young, and 4) non-reproductive females in the group; and second, giraffe form matrilineal societies, evidenced by 1) male dispersal, 2) female philopatry, 3) assistance in raising or protecting offspring, and 4) individual benefits gained from social foraging.
3. We reviewed 404 papers on giraffe behaviour and social organisation; captive studies were included where they supplemented information from free-living populations.
4. We show that giraffe exhibit many of the features typical of mammals with complex cooperative social systems and matrilineal societies. However, the social complexity hypothesis posits that such species also require complex communication systems to regulate interactions and relations among group members; giraffe communication systems are poorly understood.
5. Quantifying the fitness and survival benefits of the giraffe’s social organisation is necessary to ensure its long-term survival. Giraffe numbers have declined by 40% since 1985, they have been declared extinct in seven (possibly nine) countries and are listed as Vulnerable by the International Union for Conservation of Nature. We identify research areas that will advance our understanding of giraffe behaviour and conservation requirements.

     

The other facets of family life and their role in the evolution of animal sociality

Family life forms an integral part of the life history of species across the animal kingdom and plays a crucial role in the evolution of animal sociality. Our current understanding of family life, however, is almost exclusively based on studies that (i) focus on parental care and associated family interactions (such as those arising from sibling rivalry and parent‐offspring conflict), and (ii) investigate these phenomena in the advanced family systems of mammals, birds, and eusocial insects. Here, we argue that these historical biases have fostered the neglect of key processes shaping social life in ancestral family systems, and thus profoundly hamper our understanding of the (early) evolution of family life. Based on a comprehensive survey of the literature, we first illustrate that the strong focus on parental care in advanced social systems has deflected scrutiny of other important social processes such as sibling cooperation, parent–offspring competition and offspring assistance. We then show that accounting for these neglected processes – and their changing role over time – could profoundly alter our understanding of the origin and subsequent evolution of family life. Finally, we outline how this ‘diachronic’ perspective on the evolution of family living provides novel insights into general processes driving the evolution of animal sociality. Overall, we infer that the explicit consideration of thus‐far neglected facets of family life, together with their study across the whole diversity of family systems, are crucial to advance our understanding of the processes that shape the evolution of social life.     

Trait flexibility of generalist invertebrates exposed to contrasting predation and drying stressors

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Sacred materialism: Things and relations in a US Pagan community

Ever since Marcel Mauss wrote The Gift, the notion that things exchanged as gifts take a personalised social form, and that they partake of and mediate relational sociality, has held an important place in anthropology. Examining material practices among contemporary Pagans in the Reclaiming tradition, this article shows how they seek to imbue things with personhood, exploring the sacredness of things as alive, active and participating in social relations. In doing so, these Pagans work to forge an alternative economy founded on gift exchange and generous labour, which they hope might form a basis for a different kind of sociality from the capitalist system dominant in the United States. In practice, the encounter between gift and commodity forms in this community is a source of both conflict and emergent forms of sociality, as these alternative economic practices are fashioned around the margins of mainstream economic life.     

Evidence of interaction between Sirex noctilio and other species inhabiting the bole of Pinus

• 1 Sirex noctilio is a woodwasp native to Eurasia and Northern Africa and has recently been found infesting pines in eastern North America. Its pest status in this new range is not yet known, although it is an important pest in other areas where it has been introduced. Pinus spp. in North America are hosts to several native and alien species of subcortical insects. Interactions between the woodwasp and these species may influence its distribution or the characteristics of its life history, thus affecting its population dynamics over time.
• 2 Sixty S. noctilio‐infested Pinus spp. were felled in Ontario, Canada, and all phloem‐feeding and woodboring insects were collected and identified from each 1‐m section of the tree.
• 3 Sirex noctilio was in a tree alone 10% of the time but commonly shared the tree with subcortical beetles, such as Tomicus piniperda, Pissodes nemorensis, Ips grandicollis, Gnathotrichus materiarius and Monochamus carolinensis. The woodwasp was distributed throughout the tree stem and this distribution overlapped with that of the beetles.
• 4 Fewer but larger S. noctilio males emerged from trees with beetles compared with those without, although there was no statistical difference in females.
• 5 These findings suggest that co‐habiting beetles could negatively affect S. noctilio population dynamics over time. The potential mechanisms for this interaction are discussed.

     

Multiple environmental stressors confine the ecological niche of the rotifer Cephalodella acidophila

Summary

• 1 The planktonic food web in extremely acidic mining lakes is restricted to a few species that are either acidophilic or acidotolerant. Common metazoans inhabiting acidic mining lakes with a pH below 3 include rotifers in the genera Cephalodella and Elosa.
• 2 The life history response of Cephalodella acidophila to three environmental key factors, pH (2, 3.5, 5.0 and 7.0), temperature (10, 17.5 and 25 °C) and food concentration (10 000, 35 000 and 50 000 algal cells per mL), was investigated in a full factorial design using life‐table experiments.
• 3 The effect of each of the three environmental variables investigated on the rotifer life cycle parameters (life span, fecundity and population growth rate) differed. C. acidophila is a stenoecious species with a pH optimum in the range 3–4 and a comparably high food threshold. Combining the laboratory results with field data, we conclude that C. acidophila is severely growth limited in its natural habitat. However, low pH alone is not harmful as long as temperatures are moderate to warm and food is abundant.
• 4 The population of C. acidophila in the field is maintained mainly due to release from competitors and predators.

     

Ecological generalism facilitates the evolution of sociality in snapping shrimps

Evidence from insects and vertebrates suggests that cooperation may have enabled species to expand their niches, becoming ecological generalists and dominating the ecosystems in which they occur. Consistent with this idea, eusocial species of sponge‐dwelling Synalpheus shrimps from Belize are ecological generalists with a broader host breadth and higher abundance than non‐eusocial species. We evaluate whether sociality promotes ecological generalism (social conquest hypothesis) or whether ecological generalism facilitates the transition to sociality (social transition hypothesis) in 38 Synalpheus shrimp species. We find that sociality evolves primarily from host generalists, and almost exclusively so for transitions to eusociality. Additionally, sponge volume is more important for explaining social transitions towards communal breeding than to eusociality, suggesting that different ecological factors may influence the independent evolutionary origins of sociality in Synalpheus shrimps. Ultimately, our results are consistent with the social transition hypothesis and the idea that ecological generalism facilitates the transition to sociality.     

A nectar-secreting gall wasp and ant mutualism: selection and counter-selection shaping gall wasp phenology, fecundity and persistence

Abstract.

• 1 The natural history of a gall wasp including interactions with inquilines, parasites, and a mutualistic ant are examined. The stability of the system is described from the perspective of influences on gall wasp life history characteristics.
• 2 An exclusion experiment demonstrated that the nectar-secreting gall of Disholcaspis perniciosa mediates a mutualism with the tending ant, Formica obscuripes. Survivorship increased from 0% in the absence of ants to 25.3% in their presence, largely due to the exclusion of inquilines.
• 3 Specialized parasites, Eudecatoma spp., attacked before the ant-gall interaction began, when the developing gall was still beneath the host plant (Quercus gambellii) epidermis and ants were not in attendance. They may select for later developing gall wasps, which benefit by having fewer individuals parasitized. However, counter-selection for earlier development may result from decreased gall wasp size, decreased fecundity, and an increase in gall failures resulting from late development.
• 4 Local persistence of the gall wasp population despite increased pressure from inquilines and parasites was attributed to gall wasp escape in time due to polymorphic emergence resulting from diapause. Most individuals emerge at the end of the summer, but approximately 15% remain in the galls as prepupae for 1–5 years.

     

Impact of Social Ties on Dispersal,Reproduction and Dominance in Feral House Mice (Mus musculus domesticus)

The existence of a relationship between the social ties an individual has to other family members and its further role within the family was tested in feral house mice (Mus musculus domesticus), according to the ‘Social cohesions hypothesis’. It is predicted by the hypothesis that individuals not forming strong social ties are the first who emigrate. House mice were studied using a population cage system that allowed continuous observation of individually marked animals. Data on time staying with other animals (social ties), aggressive interactions, body weight, reproduction, and emigration were collected daily. The results may be summarized as follows:

• 1 Male emigrants were less integrated in cohorts of male littermates compared with their brothers of the same age. These male cohorts appeared to protect single males from attacks by the dominant male. No difference could be observed in social ties to other family members.
• 2 After weaning, there was no difference in social ties of male and female offspring. However, after sexual maturation social ties of males decreased significantly while those of females remained almost constant.
• 3 Female emigrants showed the same intensity of social ties as their resident sisters.
• 4 No difference could be found between social ties of females becoming pregnant and their nonreproductive sisters of the same age. Reproduction or reproductive suppression could not be explained by having more or less contact with other reproductive females.
• 5 Dominant males spent least of all time with other family members.

The social cohesions hypothesis has to be rejected in analysing proximate causes of emigration. In house mice, male emigration was caused by aggression of the dominant male in competition for the top rank within the group. This was enhanced by a lower integration in the group of same-aged brothers but is not related to a lack of integration into the family.