A Model of Mutual Tolerance and the Origin of Communal Associations Between Unrelated Females

We use a genetic algorithm model employing game theory to explore the ecological conditions favoring reproductive tolerance between two unrelated females that meet at a nesting site (i.e., breeding resource). Each female adopts one of three strategies: (1) fight for exclusive use of the nest, (2) tolerate the other female and breed communally, but fight back if attacked, or (3) leave in search of new breeding opportunities. Nests vary in the number of offspring they can support and their probability of failure due to discovery by competitors. The model predicts that communal associations are most likely to arise when (1) the benefits of nest sharing to females exceed the losses to individual reproduction, (2) additional nesting sites are rare, (3) females have limited clutch sizes, and (4) dominant females are able to skew reproduction in their favor. The amount of reproductive skew a dominant (larger) female can acquire while maintaining a communal association is predicted to increase when the asymmetry in fighting ability between females increases, and at nests that have a relatively high probability of nesting success for solitary females. When the losers of fights can parasitize the winner's brood, dominant females must reduce reproductive skew to promote a communal relationship. We discuss the ability of our model to predict patterns of facultative communal behavior in burying beetles (Silphidae; Nicrophorus spp.), as well as the absence of communal behavior in dung beetles (Scarabaeidae).     

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.     

木蜂的巢内行为和交配策略

木蜂Xylocopa作为蜜蜂科的一个重要类群, 用来研究昆虫社会进化早期阶段具有重要意义。本文综述了近年来木蜂的营巢习性、 社会性行为和交配策略的研究进展。木蜂巢的建筑与巢内巢室的分布、 巢室的大小、 贮备蜂粮的效率和被寄生的敏感性等均有一定的关系。在筑巢地点, 随着木蜂种群数量的增加, 同种雌蜂之间的资源存在着竞争, 木蜂对巢室的守卫加强, 更多地表现为对同种雌蜂的守卫以及防御其他的天敌入侵。其社会多态性由独栖性向社会性演化, 主要表现为独栖性、 亚社会性、 半社会性、 共生性和准社会性等方式, 甚至同一种群内会出现不同的社会性行为。在交配策略上, 木蜂的雄蜂在外部形态特征上具有适应交配和寻找雌蜂的进化特征, 并且具有复杂的领地行为。这些研究理论对于我们深刻理解木蜂的行为生物学具有重要意义, 同时也有助于进行更深层次的社会性蜂类演化进程的探索。     

Why do female lizards lay their eggs in communal nests?

1. In many reptile species, females oviposit communally (i.e. many clutches are laid within the same nest). This behaviour might result from constraint (scarcity of nest-sites offering suitable incubation conditions) or adaptation (direct fitness benefits accruing from the proximity of other eggs). 2. To test between these alternatives, we gathered field and laboratory data on montane scincid lizards Bassiana duperreyi from south-eastern Australia. Our data support the adaptationist hypothesis. 3. In the field, communal vs. solitary clutches were laid in similar sites, and the relative frequency of communal nesting was not predictable from nest-site availability. Thermal regimes for incubation did not differ between communal vs. solitary nests, nor between eggs at the core vs. periphery of a communal nest. In the laboratory, females selectively oviposited beside existing eggs rather than in otherwise identical potential nesting sites. 4. From cycling-temperature incubation in the laboratory, eggs incubated within a cluster of other eggs took up less water, but produced hatchlings that were larger and faster-running than were hatchlings from eggs incubated alone. 5. Hydric modifications of incubation conditions within a cluster of tightly packed eggs thus may provide a direct fitness benefit to communal oviposition.     

Microsatellites provide insight into contrasting mating patterns in arribada vs. non-arribada olive ridley sea turtle rookeries

Molecular studies of sea turtles have shown that the frequency of multiple paternity (MP) varies between species, and between rookeries of the same species. This study uses nuclear microsatellite markers to compare the incidence of MP in two neighbouring olive ridley rookeries on the Pacific coast of Costa Rica, with contrasting nesting behaviours -- the 'arribada' population nesting at Ostional and the solitary nesters of Playa Hermosa. Using two highly polymorphic microsatellite markers, we tested 13 nests from each location and found a significant difference (P < 0.001) between the level of MP of the arribada rookery (92%- the highest found for marine turtles) and that of the solitary nesting rookery (30%). Additional analyses based on six microsatellite loci revealed no genetic differentiation between nesting females from the two locations, or between nesting females and attendant males from the Ostional breeding area. Sixty-nine per cent of the nests with MP were fathered by a minimum of three different males, and three nests showed evidence of at least four fathers. The results suggest that the differences observed in levels of MP between arribada and solitary rookeries are due to an effect of abundance of individuals on the mating system. This is supported by a regression analysis combining other paternity studies on sea turtles which shows that levels of MP increase with increasing abundance of nesting females.     

Living with strangers: direct benefits favour non-kin cooperation in a communally nesting bird

The greater ani (Crotophaga major), a Neotropical cuckoo, exhibits an unusual breeding system in which several socially monogamous pairs lay eggs in a single nest and contribute care to the communal clutch. Cooperative nesting is costly-females compete for reproduction by ejecting each other's eggs-but the potential direct or indirect fitness benefits that might accrue to group members have not been identified. In this study, I used molecular genotyping to quantify patterns of genetic relatedness and individual reproductive success within social groups in a single colour-banded population. Microsatellite analysis of 122 individuals in 49 groups revealed that group members are not genetic relatives. Group size was strongly correlated with individual reproductive success: solitary pairs were extremely rare and never successful, and nests attended by two pairs were significantly more likely to be depredated than were nests attended by three pairs. Egg loss, a consequence of reproductive competition, was greater in large groups and disproportionately affected females that initiated laying. However, early-laying females compensated for egg losses by laying larger clutches, and female group members switched positions in the laying order across nesting attempts. The greater ani, therefore, appears to be one of the few species in which cooperative breeding among unrelated individuals is favoured by direct, shared benefits that outweigh the substantial costs of reproductive competition.     

Antifungal skin bacteria,embryonic survival,and communal nesting in four-toed salamanders, <Emphasis Type="Italic">Hemidactylium scutatum</Emphasis>

We examined a novel hypothesis for the maintenance of communal nesting in the salamander, Hemidactylium scutatum, namely that communal nests are more likely than solitary nests to be associated with cutaneous antifungal bacteria, which can inhibit fungal infections of embryos. A communal nest contains eggs of two or more females of the same species. The nesting behavior of H. scutatum females and survival of embryos were determined by frequent nest surveys at three ponds. For communal nests, embryonic survival tended to be higher and catastrophic nest failure was lower. Pure bacterial cultures of resident species were obtained from the salamanders’ skins by swabbing and tested against a fungal pathogen of embryos (Mariannaea sp.) in laboratory assays. We found that 27% of females had skin bacteria inhibitory to Mariannaea sp. Communal nests were more likely to have at least one female with antifungal bacteria than were solitary nests. Using a culture-independent assay (denaturing gradient gel electrophoresis of 16S rRNA gene fragments), we found that bacterial species on females and embryos were more similar to each other than they were to bacterial species found in soil within the nest, suggesting that females transmitted skin bacteria to embryos. The presence of anti-Mariannaea skin bacteria identified from the laboratory assays did not prevent fungal presence in field nests. However, once a nest was visibly infected with fungi, presence of anti-Mariannaea bacteria was positively correlated with survival of embryos. Microbe transmission is usually thought to be a cost of group living, but communal nesting in H. scutatum may facilitate the transmission of antifungal bacteria to embryos.     

An ecological analysis of mating biology of Xylocopa virginica in southern Ontario

1. Xylocopa virginica virginica Linnaeus is a wide‐ranging species with plastic nesting behaviour that appears to represent an intermediary between solitary and social nesting species. Over 3 years, a natural population was studied with the objective of quantifying the relationship among population dynamics, climate, female nest provisioning behaviour, and male mating strategy. 2. Males in the population congregated around female‐occupied nesting sites before the beginning of nest provisioning by females; both resident and satellite male mating strategies were observed. Overall, the present results are consistent with female defence polygyny. 3. Male mating strategies were consistent across the three breeding seasons of our study, in spite of annual variation in population size, sex ratio, and weather. Male mating behaviour was also consistent with that seen in other populations with longer breeding seasons. 4. Adult non‐breeding females that never leave nests are observed in nests throughout the breeding season and we hypothesise that males continue to defend territories after breeding females have mated because of a small probability they can mate with one of these non‐breeding females. 5. These results are important to our understanding of the relationship between mating systems and the evolution of sociality, contributing data on the role of ecological factors to male mating behaviour. Collection of such data for a variety of species that differ in sociality is necessary for the comparative analysis that is required to fully elucidate coevolution of mating systems and sociality.     

Solitary and group nesting in the orchid bee <Emphasis Type="Italic">Euglossa hyacinthina</Emphasis> (Hymenoptera,Apidae)

Summary Orchid bees (Euglossini) provide a potentially informative contrast for examining origins of advanced social behaviour in bees because they are the only tribe in the apine clade that do not form large colonies or have queens and workers. We investigated natural nests of Euglossa hyacinthina Dressler, an orchid bee that nests singly or in groups. By comparing the two types of nests, we examined if individuals in a group merely share the nest (are communal) or exhibit a level of social organization where there is reproductive division of labour among the females. Observations are consistent with communal nesting, indicating that all females in group nests are reproductively similar to the solitary nesting females because the provisioning of young, as well as the ovary development and mating status of females sharing nests were not different than that of solitary-nesting females. Also, multiple female nests did not produce a female-biased brood as predicted for nests with reproductive division of labour. We also investigated potential advantages of group nesting vs. individual nesting. We demonstrate that per capita offspring production is lower in nests with more than one female. However, we found that nests with single females were left unattended for longer periods of time during foraging, and that there was a high incidence of natural enemy attack in nests when females were absent. Group and solitary nesting may be advantageous under different conditions.Received 3 December 2002; revised 7 March 2003; accepted 2 April 2003.     

An experimental analysis of joint nesting in the salamander Hemidaetylium scutatum (Caudata: Plethodontidae): the effects of population density

Three nesting behaviour patterns are documented in the plethodontid salamander Hemidactylium scutatum. A female may lay eggs (1) in a solitary nest and brood them, (2) in a joint nest and brood them as well as eggs of other females, or (3) in a joint nest that is brooded by another female. The hypothesis that population density was positively associated with joint nesting was tested by following two populations for 5 years and by experimentally manipulating the population density of nesting females in artificial habitats for the latter 2 years. The proportion of joint nests did not vary with density, although joint nests tended to contain eggs of more females at the high population density. Joint nests were usually brooded by one female; thus, most females that laid eggs in joint nests did not brood them at high density. The reproductive success, as measured by survival of embryos, of solitary and joint nesters was equivalent. Joint nests were deserted less often, however, which decreased the probability of catastrophic mortality. The number of days of brooding was significantly positively correlated with loss of body mass of females, suggesting a cost to brooding behaviour. Joint nesting with solitary brooding is not explained by aggressive usurpation of nests or by brood parasitism.     

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.     

Burrow sharing and nest transfer in the digger wasp Philanthus gibbosus (Fabricius)

Philanthus gibbosus is a solitary, ground-nesting wasp in which several individuals have been observed to occupy a single burrow. Evidence is presented which indicates that several emerging males and females often occupy the parental nest for several days, the males sometimes for life. The females, however, soon disperse to found new nests, although one may continue to occupy and expand the parental nest. It appears that the larger female often maintains possession of the nest, although the interactions have not been observed. On rare occasions two females may provision a single nest for a brief period. P. gibbosus may represent a unique stage in presociality in which temporary communal nesting occurs at certain phases of the nesting cycle.     

Reproductive success of solitarily and communally nesting white-footed mice and deer mice

To determine the fitness consequences of communal nesting inwhite-footed mice, Peromyscus leucopus, and deer mice, P. maniculalus,I compared the reproductive success of field populations offemales nesting solitarily, in communal groups of more thanone female, in extended families of successive litters, andin communal groups with extended families. Mean first littersize of weanlings and juveniles 6 weeks old did not differ significantlyfor pups raised under the four nesting situations. Similarly,for pups born into extended families, litter sizes of pups fromsecond litters did not differ significantly from those of firstlitters or from pups born to solitarily nesting females. Delayeddispersal of juvenile females did not result in resource competitionor inhibition of reproduction. Thus, reproductive success offemales was not significantly affected by additional membersin the nest. At least 26 of 28 communally nesting females wereclose relatives. Solitary nesting is the common breeding patternin Peromyscus, and extended families and communal nesting arealternative reproductive tactics in response to limited space,delayed dispersal, and local grouping among related females.     

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.     

Costs of large communal clutches for male and female Greater Rheas Rhea americana

The breeding system of the Greater Rhea Rhea americana is almost unique among birds as it combines harem polygyny and sequential polyandry, with communal egg-laying and uniparental male care. In this species, large communal clutches (more than 30 eggs) are rare and have a lower hatching success than smaller clutches. Here we analyse the proximate causes of hatching failures and the costs of large communal clutches (and therefore the costs of extensive polygyny) for males and females. We evaluated if length of the nesting period, egg viability, egg losses during incubation and male parental activity at the nest were affected by clutch size. We also evaluated if chicks hatched from large clutches have a lower survival during the first 2 months after hatching. Large clutches had longer nesting period and lower hatching success, mainly as a result of bacterial contamination of the eggs and increased hatching asynchrony. In addition, large clutches tended to lose more eggs as a result of accidental breakage or predation. Male activity at the nest and chick survival were not related to clutch size. Low hatching success, nest predation risk and energetic costs associated with large clutches penalize females that join large harems and males that accept additional eggs into the nest.     

Do female degus communally nest and nurse their pups?

We examined the nesting and nursing behavior of females of the caviomorph rodent Octodon degus. We recorded the behavior of two, three-female groups of lactating degus kept in captivity and compared it with that of singly housed lactating females. Grouped females spontaneously nested communally. Five of six lactating females had non-offspring pups hanging from their teats, which suggests that some non-offspring nursing took place. Non-mothers increased their maternal behavior after the delivery of their own pups. Although time allocated to parental care by each communally nesting dam did not differ from that of solitary lactating females, young of communal litters spent less time alone and enjoyed more time cared for by a lactating female than young of single dams. Electronic Publication     

Biology of <Emphasis Type="Italic">Lasioglossum</Emphasis> (<Emphasis Type="Italic">L.</Emphasis>) <Emphasis Type="Italic">majus</Emphasis> (Hymenoptera: Halictidae), a largely solitary sweat bee with behavioural adaptations to communality

Within the bee family Halictidae there have been three origins of sociality. Although detection of origins and reversal from sociality requires phylogenetic studies, at a behavioural level a predisposition to sociality can be detected by analysis of intra-specific interactions. We studied aspects of nesting biology and behavioural interactions in Lasioglossum (Lasioglossum) majus, a poorly known halictine inhabiting temperate regions of Europe, which is suspected to be solitary. Nests were found to be largely used by one female, but some were shared by more than one individual. These few nests, whose entrances were very close to each other, were found to be connected underground. A few individuals were observed to enter in a nest where a female was waiting, behaving as a guard and allowing the incoming bee to enter the nest. By use of circle-tube experiments, the behavioural repertoire exhibited by females during encounters was assessed. Levels of withdrawal and cooperative events were comparable with those observed in other solitary nesting species, but aggressive events were very rare, as in several observed communal species. We conclude that L. (L.) majus females, despite general solitary nesting, possess behavioural components enabling them to adopt, probably in high nest-density areas, nest-sharing strategies. A similar kind of local social polymorphism has been observed in two other species of the subgenus Lasioglossum, but these are the first data available on a European species and the first record of subterranean connections among nests of halictid bees.     

Movements of Eiders Somateria mollissima on the east coast of Britain

Movement patterns, sex differences in natal dispersal and breeding dispersal, and interchange of birds between colonies were studied in the population of Eiders Somateria mollissima breeding on the east coast of Britain. First-winter Eiders reared at the Sands of Forvie, Grampian, remain at or close to the colony, while most adults move about 100 km south to winter on the Firths of Forth and Tay. A proportion of the Forvie population is sedentary. Eiders which breed in Northumberland either move north to winter on the Tay and Forth estuaries or remain close to their breeding areas. Eiders breeding in Fife are sedentary. Recoveries of British-ringed Eiders in Scandinavia indicate that some British-born males join the Baltic breeding population, probably by pairing with Scandinavian females wintering in Britain. There is extensive natal dispersal of males from Forvie, with more than twice as many Forvie-bred females as males returning to the colony to breed. The breeding dispersal of males is also twice that of females. Dispersal of males from the relatively sedentary Forvie wintering population is less than that from the breeding population. Previous work suggested that at Forvie sedentary birds nesting close to the estuary were genetically isolated from migratory ones nesting along the coast. This situation is less clear cut than had been supposed previously, with many migrants nesting close to the estuary. It is unlikely that the genetic differences between females nesting in different parts of the Forvie colony will remain stable in the long term, due to the natal and breeding dispersal of males.     

Nest re-use and communal nesting inMicrothurge corumbae (Hymenoptera,Megachilidae), with special reference to nest defense

Summary Observations on the nesting activities ofMicrothurge corumbae, carried out at the University Campus of Ribeirão Preto, São Paulo, Brazil, from 1977 to 1981, indicated that 61.9 % of nests were re-used by succeeding generations. Re-use by one generation was more frequent than by two generations, and re-use by a third was observed only once. Nests were re-used by one or several females. Single females were more frequently in the first re-use. In these cases nest re-use did not differ essentially from the solitary foundation of a new nest, except for the adoption of a pre-existing nest without excavation. In multifemale nests, analysis of relative age (wing wear), ovarian and spermathecal conditions of associated females and the content of nests at excavation indicated that the social pattern in such colonies is communal. There is some evidence that the associated females are relatives. The chalcidoid waspLeucospis was the principal nest parasite, and ants of the genusCrematogaster were nest predators. In multifemale nests, the rate of parasitism was significantly lower than in solitary nests, indicating that nest-sharing resulted in improved nest defense. On the other hand, the absence of predation on immatures of the first generation of M.commbae in multifemale nests suggests that such nests are also more resistant to attack by predators.     

An alternative route to coloniality in the bearded tit: females pursue extra-pair fertilizations

We combine behavioral studies with DNA-fingerprinting of a populationof bearded tits Panurus biarmicus and test the idea that thepursuit of extra-pair copulations by females acts as selectivepressure that may contribute to the formation of colonies inthis socially monogamous species. We show that the rate of extra-pairpaternity in colonial breeding pairs is very high, whereas noextra-pair chicks were detected in solitary nests. We foundno cases of extra-pair maternity and intraspecific brood parasitismin solitary nests, whereas it did occur in dense breeding situations.We further found no difference in the quality of males betweencolonial and solitary individuals. However, colonial femaleswere of significantly higher quality in terms of body size andcondition. We propose that high-quality females settle in coloniesto increase their opportunities to adjust the choice of theirsocial partner by obtaining extra-pair copulations. We suggestthat colony formation enables high-quality females to incitemale-male competition for extra-pair copulations and consequentlysecure extra-pair fertilizations by high-quality males and thathigh-density nesting is controlled by female bearded tits