Spatial distribution of communal nests in a colonial breeding bird: benefits without costs?

Abstract The spatial organization of individuals, or groups of individuals, within a population can provide valuable information about social organization and population dynamics. We analysed the spatial distribution of nests of the sociable weaver (Philetairus socius) on two farms in the Kalahari. Sociable weavers build large communal nests on big savannah trees, forming a pattern of trees with and without nests. We used two spatial statistics, Ripley's K and the pair correlation function, to describe characteristics of the point patterns over a range of distances. (i) At distances of 200 and 300 m, communal nests were clustered. (ii) At distances greater than 1000 m, communal nests were regularly distributed. These findings are independent of the spatial distribution of trees. Furthermore, we used Moran's I to analyse spatial autocorrelation of nest sizes. We expected negative autocorrelation because of competition between nests. But on both farms there was no significant autocorrelation of nest sizes for any distance class. The regular distribution observed at larger distances may indicate competition and/or territoriality among different nests, but the lack of spatial autocorrelation between nest sizes suggests that these interactions may happen between nest clusters rather than between single nests. This was confirmed by significant clustering of nests on small scales. We thus suggest, that colonies of P. socius consist of several nests on adjacent trees forming a cluster of subcolonies. The question why sociable weavers establish subcolonies instead of adding more chambers to the natal nest, could not simply be answered by limitation of nesting space. We suggest a strategy to avoid costs due to increasing colony size.     

Communal roosting,thermoregulatory benefits and breeding group size predictability in cooperatively breeding sociable weavers

Extreme temperatures impose energy costs on endotherms through thermoregulation and different adaptations help individuals to cope with these conditions. In social species, communal roosting and huddling are thought to decrease the energetic requirement of thermoregulation under low temperatures. This is likely to represent an important mechanism by which individuals save energy during the coldest parts of the year and hence to represent a non‐breeding benefit of sociality. Here, we investigate the potential thermoregulatory benefits of group living in roosting groups of sociable weavers Philetairus socius, a colonial cooperatively breeding passerine that builds communally a massive nest structure with several independent chambers wherein individuals breed and roost throughout the year. To investigate the benefits of sociality during the non‐breeding season, we studied the thermal environment during roosting in relation to group size. In addition, to understand the link between non‐breeding and breeding sociality in this species we studied group size stability between the pre‐breeding and breeding periods. As expected, we found that the nest chamber's night‐time temperature is strongly related to the number of birds roosting together, especially during cold nights. Specifically, birds in larger groups spent less time below the critical thermal minimum temperature (i.e. the temperature below which energy expenditure increases substantially). They were less exposed to external temperature variations. We also found a positive relationship between the number of birds roosting during winter and the breeding group size, indicating breeding group size predictability. In cooperative breeders such as the sociable weaver, the costs and benefits of sociality are usually studied during the breeding period. This study shows that a better understanding of non‐breeding benefits of group membership and group dynamics between the non‐breeding and breeding periods are necessary for a comprehensive understanding of the benefits of sociality.     

Sociable Weavers Increase Cooperative Nest Construction after Suffering Aggression

The major transitions in evolution rely on the formation of stable groups that are composed of previously independent units, and the stability of these groups requires both cooperation and reduced conflict. Conflict over group resources may be common, as suggested by work in both cichlids and humans that has investigated how societies resolve conflict regarding investment in group resources, i.e. public goods. We investigated whether sociable weavers (Philetairus socius) use aggressive behaviors to modulate the cooperative behavior of group mates. We find that the individuals that build the communal thatch of the nest, i.e. the individuals most at risk of exploitation, are the most aggressive individuals. We show that individuals that invest in interior chamber maintenance, possibly a more selfish behavior, suffer relatively more aggression. After suffering aggression individuals significantly increase cooperative construction of the communal nest thatch. We show that cooperative individuals target aggression towards selfish individuals, and the individuals suffering aggression perform cooperative behaviors subsequent to suffering aggression. In addition to other evolutionary mechanisms, these results suggest that aggression, possibly via the pay-to-stay mechanism, is possibly being used to maintain a public good.     

Cooperative investment in public goods is kin directed in communal nests of social birds

The tragedy of the commons predicts social collapse when public goods are jointly exploited by individuals attempting to maximize their fitness at the expense of other social group members. However, animal societies have evolved many times despite this vulnerability to exploitation by selfish individuals. Kin selection offers a solution to this social dilemma, but in large social groups mean relatedness is often low. Sociable weavers (Philetairus socius) live in large colonies that share the benefits of a massive communal nest, which requires individual investment for construction and maintenance. Here, we show that despite low mean kinship within colonies, relatives are spatially and socially clustered and that nest‐building males have higher local relatedness to other colony members than do non‐building males. Alternative hypotheses received little support, so we conclude that the benefits of the public good are shared with kin and that cooperative investment is, despite the large size and low relatedness of these communities, kin directed.     

THE THERMAL SIGNIFICANCE OF THE NEST OF THE SOCIABLE WEAVER PHILETAIRUS SOCIUS: SUMMER OBSERVATIONS

Extending a previous study of the thermal significance of the nest of the Sociable Weaver carried out during the winter, we measured temperature and humidity in the matrix and chambers of a large nest of this species in the Kalahari Gemsbok National Park, South Africa, during the austral summer of December 1973. Air temperatures outside the nest ranged from 16 to 33.5°C but temperatures in occupied chambers varied over a range of only 7 or 8°C and remained well within the zone of thermal neutrality for a passerine bird of this size. Compared to outside air temperatures, those within the nest matrix were lower during the day and higher at night. Thus, the nest ameliorates the effects of external temperatures and allows maintenance inside the chambers of a range of temperature favourable to the birds. In winter we found up to five roosting adults per chamber, with some chambers left empty. In the same nest in summer we found no more than two adults per chamber but virtually all chambers were occupied. The principal mechanism for maintaining chambers within the zone of minimal energetic cost is changes in the number of birds in the nest chambers at night. Humidity inside the occupied and unoccupied chambers was somewhat higher in the former but always less than that of outside air in both cases. Air movement through the desiccated nest materials causes uptake by these materials of most of the water vapour introduced by the birds, and this moisture is dissipated to the outside during the day so that the nest remains dry. The highly social and colonial habits of the birds and their year-round occupancy and maintenance of the nest favour a system of opportunistic breeding that may be initiated by rainfall at any season. Larger nests provide the most favourable environment for energy conservation and successful reproduction. Even the largest nests, however, do not prevent predation during the warm season by snakes such as the Cape Cobra, which may consume all the eggs and young in all the chambers of a large nest. The effects of such heavy predation may be offset by the birds' capability for breeding during times too cold for reptile activity. It seems likely that in smaller nests such as those on telephone poles, lack of predation would favour summer breeding while thermal problems would limit breeding success in winter. In larger nests, breeding success may be lower in summer because of predation and higher in winter when reptile predation is lacking and thermal problems are minimized by the nest structure. The large nest not only makes possible the success of the Sociable Weaver in desert areas, but the nest could only exist in such areas and the species' range is thereby restricted. Higher humidity and heavier rainfall would cause fermentation within the nest mass, loss of its thermoregulatory advantages, and ultimately its decomposition and destruction. Therefore, the unique nesting system of the Sociable Weaver appears to be initially self-reinforcing and ultimately self-limiting.     

To nest communally or not to nest communally: a review of rodent communal nesting and nursing

Communal nesting, the sharing of parental responsibilities between multiple individuals in a nest, is common in many rodents. Upon first glance, this behaviour seems to be selectively disadvantageous. Communal care not only involves energetic costs, but may also be subject to cheating behaviour. Despite abundant literature, many questions remain regarding advantages gained by females that form nesting groups. I discuss the communal nesting of eusocial, singular and plural breeding rodents. I first clarify the distinction between communal nesting and thermoregulatory huddling. I then review two major groups of hypotheses ('ecological constraints' and 'benefits of philopatry') that are used to explain the occurrence of communal nesting in rodents. Most likely, these hypotheses are not mutually exclusive. Regardless of the main selective pressure(s) acting on communal nesting, the indirect components of inclusive fitness that result from nest sharing most likely influence the formation and maintenance of communal nests. Communal nesting and nursing (the sharing of milk with nonoffspring) are under different selective regimes and thus, must be evaluated separately. I review adaptive and nonadaptive hypotheses for rodent communal nursing. I argue that communal nursing may not be adaptive as mothers may be forced to share milk with nonoffspring in large communal nests (i.e. communal nursing may be a cost associated with communal nesting). In conclusion, I propose directions for future study that may improve our understanding of communal nesting and nursing in the wild. Copyright 2000 The Association for the Study of Animal Behaviour.     

At home with the birds: Kalahari tree skinks associate with sociable weaver nests despite African pygmy falcon presence

The way in which animals use habitat can affect their access to key resources or how they are buffered from environmental variables such as the extreme temperatures of deserts. One strategy of animals is to modify the environment or take advantage of structures constructed by other species. The sociable weaver bird (Philetairus socius) constructs enormous colonial nests in trees. These nests are frequented by Kalahari tree skinks (Trachylepis spilogaster) and the two species coexist over a large portion of the Kalahari Desert in South Africa. We tested whether skinks were more abundant in trees containing sociable weaver nests and asked whether the physical features of trees were important predictors of skink abundance. We then focused on potential costs of this association by examining the relationship between skink abundance and the presence of a potential predator, the pygmy falcon (Polihierax semitorquatus), which exclusively uses weaver nest colonies for roosting and nesting. Finally, we simulated a predatory threat to determine if skinks assess risk differently if a weaver nest is present. We found a significant positive association between the presence of weaver nests and skink abundance. In the absence of nests, the type of tree did not influence skink abundance. Skinks used weaver nests and were more likely to perch on the nest than the tree. When threatened with predation, skinks preferred to take refuge in nests. Surprisingly, the presence of nesting pygmy falcons in nests did not influence skink abundance, perhaps because of the abundance of nearby refuges within nests, tree crevices, or in debris at the tree base. We suggest that sociable weaver nests provide multiple benefits to skinks including lowered predation risk, thermal refuges and greater prey availability, although this requires experimental testing. In the current era of global climate change, sociable weaver nests may become a crucial resource for skinks seeking refuge as the Kalahari climate warms.     

Elaborate Nests in a Weaverbird: A Role for Female Choice?

Studies on sexual selection have focused on behaviour and morphology, but several groups of animals build elaborate structures associated with acquiring a mate. I investigated female choice for nests built by male baya weavers (Ploceus philippinus). Nest choice by females should be strong, as nests are obvious direct benefits provided by males. I used a field experiment supplemented with correlational information to ask whether females appear to base mate choice decisions on male behaviour, nest architecture, and nest location. When the nests of highly visited males were exchanged with those of poorly visited males, female visits remained highest at the original male and location. I found no relationship between female choice and male display or other behaviour. Correlational analyses show that nest location was a better predictor of female choice than was nest architecture. These data suggest that current female choice is driven more by access to safe nesting sites rather than to well‐built nests, possibly because all males are able to build nests of adequate quality. However, nest architecture is unlikely to be irrelevant to females, and its role deserves further investigation.     

Why do grebes cover their nests? Laboratory and field tests of two alternative hypotheses

Egg predation is a common feature influencing the reproductive success of open nesting birds. Evolutionary pressure therefore favours building cryptic, inconspicuous nests. However, these antipredatory pressures may be in conflict with thermoregulatory constraints, which select for dry nest material maintaining optimum temperature inside a nest cup during the absence of incubating parents. Here we examined possible trade-offs between nest crypsis and thermoregulation in Little Grebes (Tachybaptus ruficollis), which lay their eggs in floating nests built from wet plant material. As this species regularly covers its eggs with nest material, we experimentally examined (1) the rates of egg predation on covered and uncovered artificial nests and (2) possible thermoregulatory costs from nest covering by comparing temperature and relative humidity changes inside the nest cup. Results revealed that covering clutches is beneficial in terms of deterring predators, because uncovered eggs were more vulnerable to predation. Moreover, covering clutches also had thermoregulatory benefits because the mean temperature and relative humidity inside nest cups covered by dry or wet materials were significantly higher for covered compared to uncovered treatments. Covering clutches in Little Grebes therefore does not pose thermoregulatory costs.     

THE SOCIABLE WEAVER,PART 4: PREDATORS,PARASITES AND SYMBIONTS

Maclean, G. L. 1973. The Sociable Weaver, Part 4: Predators, parasites and symbionts. Ostrich 44: 241–253.

The main nest predator of the Sociable Weaver in the Kalahari sandveld is the Cape Cobra Nala nivea. This snake causes great losses of eggs and chicks; one cobra may eat the contents of an entire nest mass at one feed. Another nest predator which causes smaller losses of eggs and chicks but great destruction to the nest masses is the Honey Badger Mellivora capensis. These are the only two nest predators on the Sociable Weaver in the study area. Predators on adult Sociable Weavers include several birds of prey and some small carnivorous mammals.

Adult Sociable Weavers have few ectoparasites and hardly any Mallophaga. A common ectoparasite on the legs of chicks is a blood-sucking Dermestes larva, which appears not to be harmful. The only endoparasite found was the nematode, Diplotriaena ozouxi, which infected the abdominal air sacs of the adults.

The nest material of the Sociable Weavers' communal nest masses was inhabited by a wealth of invertebrate animals and a few harmless reptiles such as skinks and geckos.

Some of the chambers in a Sociable Weaver nest mass may be taken over by other species of birds. Most of these, such as Redheaded Finches Amadina erythrocephala, use the chambers for breeding purposes only, but the most important avian symbiont, the Pygmy Falcon Polihierax semitorquatus, is a permanent resident in the chambers. The presence of Pygmy Falcons is resented by the weavers but the falcons may help to keep snakes away from the nest mass. Adult Sociable Weavers are not normally preyed on by Pygmy Falcons, although the falcons may occasionally take young weavers in the nest chambers.

The tops of the nest masses may be used as nest sites by the Giant Eagle Owl Bubo lacteus. Barn Owls Tyto alba may use cavities in the superstructure of nest masses for roosting in. Neither of these owls appeared to prey on the weavers.     

Maternal effects in the highly communal sociable weaver may exacerbate brood reduction and prepare offspring for a competitive social environment

Maternal effects can influence offspring phenotype with short- and long-term consequences. Yet, how the social environment may influence egg composition is not well understood. Here, we investigate how laying order and social environment predict maternal effects in the sociable weaver, Philetairus socius, a species that lives in massive communal nests which may be occupied by only a few to 100+ individuals in a single nest. This range of social environments is associated with variation in a number of phenotypic and life-history traits. We investigate whether maternal effects are adjusted accordingly. We found no evidence for the prediction that females might benefit from modifying brood hierarchies through an increased deposition of androgens with laying order. Instead, females appear to exacerbate brood reduction by decreasing the costly production of yolk mass and antioxidants with laying order. Additionally, we found that this effect did not depend on colony size. Finally, in accordance with an expected increased intensity of environmental stress with increasing colony size, we found that yolk androgen concentration increased with colony size. This result suggests that females may enhance the competitive ability of offspring raised in larger colonies, possibly preparing the offspring for a competitive social environment.     

Relatedness and genetic structure in a socially polymorphic population of the spider Anelosimus studiosus

The evolution of sociality remains a challenge in evolutionary biology and a central question is whether association between kin is a critical factor favouring the evolution of cooperation. This study examines genetic structure of Anelosimus studiosus, a spider exhibiting polymorphic social behaviour. Two phenotypes have been identified: an ‘asocial’ phenotype with solitary female nests and a ‘social’ phenotype with multi‐female/communal nests. To address the questions of whether these phenotypes are differentiated populations and whether cooperative individuals are closely related, we used microsatellites to analyse individuals from both communal and solitary nests. We found no evidence of differentiation between social and solitary samples, implying high rates of interbreeding. This is consistent with the hypothesis that these phenotypes coexist as a behavioural polymorphism within populations. Pairwise relatedness coefficients were used to test whether cooperating individuals are more closely related than expected by chance. Pairwise relatedness of females sharing communal webs averaged 0.25, the level expected for half‐siblings and significantly more closely related than random pairs from the population. Solitary females collected at similar distances to the communal spider pairs were also more closely related than expected by chance (mean relatedness = 0.18), but less related than social pairs. These results imply that low dispersal contributes to increase likelihood of interaction between kin, but relatedness between social pairs is not explained by spatial structure alone. We propose that these phenotypes represent stages in the evolution of sociality, where viscous population structure creates opportunities for kin selection and cooperation is favoured under certain environmental conditions.     

Communal Nesting in Asocial Abert's Squirrels: the Role of Social Thermoregulation and Breeding Strategy

The social thermoregulation hypothesis states that endothermic species will communally nest to reduce energy expenditures on thermoregulation. The hypothesis predicts that the frequency of communal nesting should increase with decreasing ambient temperature. The potential costs of communal nesting (e.g., increased predation risk, resource competition, cuckoldry, parasite/disease transmission, or infanticide) could decrease the frequency of communal nesting especially for asocial breeding females with dependent offspring. We examined the effects of ambient temperature and seasonal reproductive activities on the probability of communal nesting in Abert's squirrels (Sciurus aberti) in the Pinaleño Mountains, Arizona. Most squirrels nested consistently with the same partner in mixed‐sex pairs. The proportion of individuals engaging in communal nesting increased with decreasing ambient temperature as predicted by the social thermoregulation hypothesis. The onset of the breeding season greatly reduced the proportion of individuals communally nesting. The negative relationship between ambient temperature and communal nesting supports the use of communal nesting in Abert's squirrels as a mechanism to reduce thermoregulatory costs during cold conditions. The abrupt drop in the frequency of communal nesting during the breeding season is likely due to female abandonment of this behavior. By avoiding communally nesting during the breeding season, females may prevent males from mating with them outside of mating chases, reduce resource competition, and protect offspring from infanticide, diseases, and parasites. Males may gain additional fitness benefits from nesting with females because familiarity with females increases dominance rank in mating activities.     

Movements,status, access to nectar,and spatial organisation of the tui

Abstract

Movements of tui (Prosthetnadera novaeseelandiae) breeding on Tiritiri Matangi Island were followed and related to changes in nectar access. The island has limited nectar sources and birds must move to gain access to nectar year round. All tui leave the island, although the extent and timing of movement varies for different individuals. Tui return for breeding but again the timing of the return varies for individuals. Nectar sources were typically concentrated in both time and space, and access was related to individual asymmetries such as sex, residential status and age. These same asymmetries also relate to the timing of individuals movements, and where individuals nest relative to nectar sources. At concentrated food sources, access was determined by an absolute social hierarchy. Although tui defend exclusive feeding territories, breeding territories are not exclusive and access to nectar is determined by distance to nest site. Such variability in spatial organisation is related to a continuum concept of social organisation.     

How does starvation affect spatial organization within nests in <Emphasis Type="Italic">Lasius niger</Emphasis>?

Spatial distribution of ant workers within the nest is a key element of the colony social organization contributing to the efficiency of task performance and division of labour. Spatial distribution must be efficiently organized when ants are highly starved and have to get food rapidly. By studying ants’ behaviour within the nest during the beginning of food recruitment, this study demonstrates how the spatial organization is affected by starvation and improves the efficiency and the speed of recruitment as well as the allocation of food. (1) In starved nests, nestmates left the deep part of the nest and crowded near the nest entrance. This modification of the spatial distribution is a local phenomenon concerning only the individuals situated in the first chamber near the nest entrance. These starved individuals have a higher probability of leaving the nest after a contact with recruiters than nestmates situated deeper in the nest. This strongly suggests that nestmates situated near the nest entrance have a low response threshold to the signals emitted by recruiters. Their higher responsiveness speeds up their exit to the foraging area and hence may increase the efficiency of highly starved colonies in exploiting new food opportunities. (2) In starved nests, the trajectory covered by recruiters between contacts with nestmates was nearly twice as small. For recruiters, this represented a gain of time in the allocation of food. As the recruitment process follows snowball dynamics, this gain of time by starved recruiters might also speed up the exploitation of food. This study evidences how the spatial distribution of individuals as a function of their motivational state might have a regulatory function in the recruitment process, which should be generic for many social species.     

Group nesting and individual variation in behavior and physiology in the orchid bee <Emphasis Type="Italic">Euglossa nigropilosa</Emphasis> Moure (Hymenoptera,Apidae)

The Euglossini are a key group for studying the traits that promote or hinder highly social behavior in bees because it is the only tribe in the Apine clade without large colonies or females with distinct life histories, e.g. queens and workers. There have been few studies on behavior of orchid bee females in nests because these nests are not found easily. Taking advantage of the relatively high abundance of Eg. nigropilosa nests at Reserva Natural La Planada, Colombia, we examined social behavior of Eg. nigropilosa individuals in five nests (3 original and 2 reused) for nine months. We report this species to have the largest colonies known for Euglossa, with nests reaching up to 22 individuals, and all nests containing more than one female bee from the same generation. These nests presented many traits that correspond to communal insect colonies. No generational overlap and no cooperative brood care were detected. We examined natural enemies and resource limitation as important factors for group nesting. We examined parasitoid attacks to cells in a nest with females and one without females. We also searched for nesting locations and examined nest re-use as indicators of nest site limitation. Lastly, we examined behavioral and physiological differences among females in the same nest. Such differences could be the bases for evolution of alternative life histories among group living females. We examined extent of ovary development and oviposition rates in similarly aged females in the same nest. We found large variation in reproductive effort of young females. We also examined differences in resin foraging and cell usurpation behaviors. Behavioral specialization was observed, with some individuals bringing only resin to the nest. Inside the nests, bees had territories in which they constructed and defended cells. This territoriality may be a defense against usurpation of provisioned cells by nest mates. Received 10 December 2007; revised 2 May 2008; accepted 7 May 2008.     

Communal nesting and communal nursing in house mice,Mus musculus domesticus

The functional significance of communal nesting and nursing is poorly understood. Female house mice often communally nest, and within these communal nests females appear to indiscriminately nurse all pups, a rare trait for any mammal. In this study, the hypothesis that communal nesting provides protection from conspecific infanticide was tested and supported in semi-natural populations of house mice. Conspecific infanticide in single-mother nests (69%, N = 412) was twice that in communal nests (33%, N = 508). Because this major benefit of communal nesting does not require communal nursing, direct benefits to communal nursing itself were tested. Most proposed benefits should result in heavier weaning weights, but no differences were found between communal and single nests in the semi-natural populations. If communal nursing is to be avoided in communal nests, dams must recognize their own pups. Retrieval tests conducted in the laboratory produced equivocal results. Dams discriminated between pups that differed in age, but not between their own and other age-matched pups. The major survival advantage of communal nesting, coupled with the failure to find nutritional advantages for communally nursed pups, supports a recent suggestion that communal nursing is an unavoidable consequence of communal nesting. This hypothesis is further strengthened by data indicating that communal nesting partners tend to be kin, thereby providing inclusive fitness benefits to communal nursing. Although costs of communal nursing were proposed and tested, no such costs were found. We also show from 15 observations of infanticide that all classes of adults (territorial and non-territorial males, pregnant and non-pregnant females) are infanticidal. These observations are in conflict with previous laboratory studies.     

Dominance and huddling behaviour in Bronze Mannikin Lonchura cucullata flocks

Bronze Mannikins Lonchura cucullata form flocks of up to 30 birds at the end of the breeding season and huddle in communal nests during winter. Assuming there are thermoregulatory benefits associated with huddling for these small endotherms, particularly during winter, we predicted that dominant individuals in flocks would obtain the central locations within the huddles. We tested this prediction by determining the dominance hierarchies of six captive flocks of five individuals by examining interactions at a feeder. Mannikins were observed to huddle when cold-stressed. Although mannikins do not naturally form linear huddles, for ease of observation we recorded the positions of individually marked birds in linear huddles at 5 °C before and after disturbances every half hour for 3 h. Captive flocks of mannikins established linear dominance hierarchies and dominant individuals obtained the central location within the huddle. The central position in the huddle may be beneficial in thermodynamically challenging environments through reduced energy expenditure.     

The Antsy Social Network: Determinants of Nest Structure and Arrangement in Asian Weaver Ants

Asian weaver ants (Oecophylla smaragdina) are arboreal ants that are known to form mutualistic complexes with their host trees. They are eusocial ants that build elaborate nests in the canopy in tropical areas. A colony comprises of multiple nests, usually on multiple trees, and the boundaries of the colony may be difficult to identify. However, they provide the ideal model for studying group living in invertebrates since there are a definite number of nests for a given substrate, the tree. Here, we briefly examine the structure of the nests and the processes involved in the construction and maintenance of these nests. We have described the spatial arrangement of weaver ant nests on trees in two distinct tropical clusters, a few hundred kilometres apart in India. Measurements were made for 13 trees with a total of 71 nests in the two field sites. We have considered a host of biotic and abiotic factors that may be crucial in determining the location of the nesting site by Asian weaver ants. Our results indicate that tree characteristics and architecture followed by leaf features help determine nest location in Asian weaver ants. While environmental factors may not be as influential to nest arrangement, they seem to be important determinants of nest structure. The parameters that may be considered in establishing the nests could be crucial in picking the evolutionary drivers for colonial living in social organisms.     

A description of nesting behaviors,including factors impacting nest site selection,in black‐and‐white ruffed lemurs (Varecia variegata)

Nest site selection is at once fundamental to reproduction and a poorly understood component of many organisms’ reproductive investment. This study investigates the nesting behaviors of black‐and‐white ruffed lemurs, Varecia variegata, a litter‐bearing primate from the southeastern rainforests of Madagascar. Using a combination of behavioral, geospatial, and demographic data, I test the hypotheses that environmental and social cues influence nest site selection and that these decisions ultimately impact maternal reproductive success. Gestating females built multiple large nests throughout their territories. Of these, females used only a fraction of the originally constructed nests, as well as several parking locations as infants aged. Nest construction was best predicted by environmental cues, including the size of the nesting tree and density of feeding trees within a 75 m radius of the nest, whereas nest use depended largely on the size and average distance to feeding trees within that same area. Microhabitat characteristics were unrelated to whether females built or used nests. Although unrelated to nest site selection, social cues, specifically the average distance to conspecifics’ nest and park sites, were related to maternal reproductive success; mothers whose litters were parked in closer proximity to others’ nests experienced higher infant survival than those whose nests were more isolated. This is likely because nesting proximity facilitated communal crèche use by neighboring females. Together, these results suggest a complex pattern of nesting behaviors that involves females strategically building nests in areas with high potential resource abundance, using nests in areas according to their realized productivity, and communally rearing infants within a network of nests distributed throughout the larger communal territory.