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.     

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。     

Comparison of social and solitary nesting carpenter bees in sympatry reveals no advantage to social nesting

Facultatively social bees allow for comparisons of social and solitary behaviour under similar environmental conditions. When such bees are polymorphic within the same population it provides a special and somewhat unusual opportunity to examine factors leading to cooperative (social) behaviours where many parameters such as environment are not variable. Some species of bees in the genus Xylocopa offers such opportunities. Studies of these bees often point to guarding and resource limitation as factors leading to social nesting. The large carpenter bee Xylocopa virginica L. (Hymenoptera: Apidae) is one species that exhibits both solitary and social nesting behaviour within the same populations. This paper first compares social and solitary nests in an Ontario population of X. virginica to determine if there is a reproductive advantage to social nesting. Following this, a series of possible explanations for social nesting, and the roles of females in social nests are examined. Social nests have similar brood sizes to solitary nests and productivity as a function of colony size is reduced with increasing number of foundresses. Additional foundresses are not effective guards, do not prevent parasitism, do not likely perform significant work, and do not assume foraging with the loss of a primary foundress. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 998–1010.     

Sociality in <Emphasis Type="Italic">Euglossa</Emphasis> (<Emphasis Type="Italic">Euglossa</Emphasis>) <Emphasis Type="Italic">viridissima</Emphasis> Friese (Hymenoptera,Apidae, Euglossini)

Euglossa viridissima is an orchid bee that forms both solitary and multiple female nests, making it a suitable species for the study of factors leading to diverse degrees of sociality in Euglossines. We conducted observations in eight reused nests (where a first generation of bees had been produced) kept in artificial boxes from the Yucatan Peninsula, Mexico. Five nests were reused (reactivated) by a single female (SFN), two nests reused by a mother and one daughter (MFN1) and one nest reused by the mother and two daughters (MFN2). No single nest was reactivated by unrelated females. The number of foraging trips, their duration and the duration of cell provisioning was not different between SFN and MFN. The overall production of cells per female was not different either between both types of nest. However, in MFN although all females did lay eggs, there was a reproductive skew in favor of the mother (95 and 45% of the brood produced in MFN1 and MFN2 respectively). She showed reproductive control of her daughters through oophagy and displaying threatening behavior when the daughters tried to open a cell where she had laid an egg. Brood losses to parasites (Anthrax sp. (Bom-byliidae) and Hoplostelis bivittata (Megachilidae)) were only found in SFN which possibly reflects and advantage of MFN in this respect. Our results coupled with other studies in Euglossa, reveal that a wide range of social behaviors occur in this genus, from solitary and communal to primitive reproductive division of labor. Multiple factors involving different levels of pressure imposed by food availability and parasites may favor such a diverse range of nesting behaviors. Interestingly, female associations in E. viridissima seem a result of kin selection that is enforced by coercion from mother females on their daughters. More studies are needed to shed light upon the social organization of Euglossa and other Euglossines and on their phylogenetic relationships in order to trace the origins of eusociality in Apidae. Received 12 February 2008; revised 25 June 2008; accepted 17 July 2008.     

An overview of proximate factors affecting the nesting behavior of solitary wasps and bees (Hymenoptera: Aculeata) in preexisting cavities in wood

Guilds of Aculeate solitary wasps and bees that nest in preexisting cavities in wood are important components of terrestrial ecosystems because they engage in several ecological interactions (e.g. predation and pollination) with other species of plants and animals. Spatial and temporal variations in richness and abundance of solitary wasps and bees can be related to changes in environmental structure and in the diversity of other groups of organisms. The nesting period of these Aculeata is their most critical life cycle stage. Females of solitary wasp and bee species invest relatively more time constructing and provisioning their nests than do females of social species. Differently from species that nest in the soil or construct exposed nests, the main factors affecting the reproductive success of solitary species nesting in preexisting wood holes are still unknown. Our objective is to provide an overview of the role of proximate causes of nesting failure or success among solitary wasps and bees (Aculeata), for designing effective conservation and management strategies for these Hymenoptera.     

Fitness consequences of ecological constraints and implications for the evolution of sociality in an incipiently social bee

Ecological constraints such as resource limitation, unfavourable weather conditions, and parasite pressure have long been considered some of the most important selective pressures for the evolution of sociality. In the present study, we assess the fitness consequences of these three ecological factors on reproductive success of solitary nests and social colonies in the socially polymorphic small carpenter bee, Ceratina australensis, based on 982 nests collected over four reproductive periods. Nest site limitation was predicted to decrease opportunities for independent nest initiation and increase the frequency of social nesting. Nest sites were not limiting in this species and the frequency of social nesting was consistent across the four brood‐rearing periods studied. Unfavourable weather was predicted to lower the frequency of female dispersal from their natal nests and to limit the brood‐rearing season; this would increase the frequency and fitness of social colonies. Daily temperature and precipitation accumulation varied between seasons but were not correlated with reproductive success in this bee. Increased parasite pressure is predicted to increase the frequency and fitness of social colonies because solitary bees must leave the nest unattended during foraging bouts and are less able to defend the nest against parasites. Severe parasitism by a chalcid wasp (Eurytoma sp.) resulted in low reproductive success and total nest failure in solitary nests. Social colonies had higher reproductive success and were never extirpated by parasites. The high frequency of solitary nests suggests that this is the optimal strategy. However, social colonies have a selective advantage over solitary nesting females during periods of extreme parasite pressure, and we suggest that social nesting represents a form of bet‐hedging against unpredictable fluctuations in parasite number. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 57–67.     

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.     

Predation Cues in Solitary bee Nests

Predation at the nesting site can significantly affect solitary bees’ reproductive success. We tested female red mason bees’ (Osmia bicornis L.) acceptance of potential nesting sites, some of which were marked with cues coming from predated conspecifics (crushed bees) or from a predator itself (rodent excreta). In our experiment, females did not avoid nests marked with either of the two predator cues. We suggest that bee females do not recognize these two cues as risky. Alternatively, costs of abandoning natal aggregation might be too high compared with any perceived predation risk of staying. Moreover, the presence of crushed bees can provide positive information about the presence of conspecifics and, possibly, information about a nesting aggregation that may be preferred by bees when choosing a nesting site.     

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.     

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).     

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.     

Evidence of Entrance Sharing and Subterranean Connections in Andrena agilissima (Hymenoptera; Andrenidae)

Andrena agilissima is a bee that nests on sloped or vertical earth walls. Data from 3 years of fieldwork presented here show that this species is communal and uses the same nesting site for many years. On an earth wall on Isola d'Elba (Tuscany, Italy), covered with many bee nest entrances, we studied an aggregation of Andrena agilissima to determine if bees share entrances, if the sharing provides evidence for subterranean connections, and if bees involved in sharing show a division of labor. We marked several females coming out of the same entrance, as well as recapturing the same females using different entrances. No division of labor was observed in our observations.     

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.     

Nesting biology of tropical solitary and social sweat bees,Lasioglossum (Dialictus) figueresi Wcislo andL. (D.) aeneiventre (Friese) (Hymenoptera: Halictidae)

Summary The nesting biology of a mainly solitary bee,Lasioglossum (Dialictus) figueresi, is compared with that of a possible relative and mainly eusocial bee,L. (D.) aeneiventre. These bees nest in the ground in highly disturbed areas in the Meseta Central of Costa Rica. Information is provided on social organization, male production, diel and seasonal activity patterns, pollen utilization, natural enemies and nest architecture. L. (D.) figueresi nests within aggregations in vertical earthen banks, and 80–90 % of females are solitary during the nest-provisioning phase. Social nests contain two (or rarely three) females which may be either equal or unequal in reproductive status (i.e. mated with developed ovaries or not). Solitary nests and two-female nests do not experience different rates of parasitism. Mid-way through the dry season, females cease provisioning at a time when otherL. (Dialictus) remain active. Females typically remain within their nests, although they occasionally forage for nectar. This behavior is similar of that of spring gynes of temperate eusocial species. The egg-to-adult developmental rate ofL. (D.) figueresi is unusually slow for halictine bees, however, so that all the adult females die before their brood eclose in April and May, precluding overlap of generations. The eclosed offspring remain in open cells within their natal nests until mid-June, when both males and females emerge to mate. These newly mated females either establish new nests or re-use old ones. L. (D.) aeneiventre nests within aggregations in horizontal ground or in vertical banks. A foundress female digs a nest at the beginning of the dry season, although some re-activate old nests. Foundress nests develop into colonies with various kinds of social organization. In contrast toL. (D.) figueresi, L. (D.) aeneiventre is active nearly all year round, except during periods of heavy rain, and produces up to three broods per year.Sweat bees (Hymenoptera: Halictinae) are a socially heterogeneous group of mainly ground-nesting bees which are abundant world-wide. Intra-specific variation in social behavior is prevalent both within and among populations, presumably indicating social and environmental control of behavioral modifications (see e.g. Sakagami and Munakata, 1972; Eickwort, 1986; Packer, 1990; Yanega, 1988; reviewed in Michener, 1990). The initial stages of hymenopteran social evolution are represented by solitary individuals and those in undifferentiated societies, yet their biology is not well known, as is true for the numerous tropical halictine species or populations (see Michener, 1990).The subgenusDialictus ofLasioglossum is a primarily New World group of several hundred species (Moure and Kurd, 1987). These bees are monotonously similar in structure and appearance, yet diverse in social behavior. FemaleLasioglossum (Dialictus) figueresi are usually solitary, and structurally are very similar to their social relativeL. (D.) aeneiventre (Wcislo, 1990 a). The systematic placement of these species with respect to otherL. (Dialictus) is uncertain, but they have no obvious affinities to other recognized species groups (G. C. Eickwort, pers. comm.). Unusual morphological features, such as large size, yellowish wings and pubescence, and features of the genital organs, may indicate thatL. (D.) figueresi is the more derived of the pair, and may therefore be secondarily solitary, as is known for other sweat bees (Packer, 1991).     

Emergence and Consequences of Division of Labor in Associations of Normally Solitary Sweat Bees

Division of labor is a pervasive feature of animal societies, but little is known about the causes or consequences of division of labor in non-eusocial cooperative groups. We tested whether division of labor self-organizes in an incipient social system: artificially induced nesting associations of the normally solitary sweat bee Lasioglossum ( Ctenonomia ) NDA-1 (Hymenoptera: Halictidae). We quantified task performance and construction output by females nesting either alone or with a conspecific. Within pairs, a division of labor repeatedly arose in which one individual specialized on excavation and pushing/tamping while her nestmate guarded the nest entrance. Task specialization could not be attributed to variation in overall activity, and the degree of behavioral differentiation was greater than would be expected due to random variation, indicating that division of labor was an emergent phenomenon generated in part by social dynamics. Excavation specialists did not incur a survival cost, in contrast to previous findings for ant foundress associations. Paired individuals performed more per capita guarding, and pairs collectively excavated deeper nests than single bees – potential early advantages of social nesting in halictine bees.     

Sociality in the phylogenetically basal allodapine bee genus Macrogalea (Apidae: Xylocopinae): implications for social evolution in the tribe Allodapini

Recent phylogenetic studies based on DNA sequence data indicate that the tropical African bee genus Macrogalea is the sister group to the remaining extant allodapine fauna, whereas previously it was thought to be a distal group. This leads to some fundamental changes in our understanding of social evolution in the allodapines. Earlier studies suggested that Macrogalea showed only weak forms of social behaviour and these were not well characterized. However, large samples of Macrogalea zanzibarica presented here show that this species exhibits marked social behaviour. Nearly half of nests collected contained two or more adult females, with up to 10 females per nest. Brood are reared progressively and brood ages within colonies are staggered, giving rise to colonies with very mixed age structures and therefore frequent opportunities for alloparental care. Ovarian dissections indicate non‐simple forms of reproductive partitioning within colonies and most multi‐female colonies show evidence that more than one female has contributed to egg production, though reproductive episodes among colony members are frequently asynchronous. Some females show signs of much higher wing wear than their nestmates, but always show signs of previous reproduction. Reproductive division of labour appears to be temporally marked, ovarian differentiation among nestmates is linked to relative body size, but permanent worker‐like or queen‐like castes appear to be absent. This is similar to the communal, continuously brooded and multivoltine behaviour of some tropical halictine species and may be due to the aseasonal nature of brood development in tropical regions. Patterns of per capita brood production indicate large benefits to multi‐female nest occupancy, and sex allocation is strongly female biased. These findings strongly suggest that the capacity for complex social interactions and alloparental care is an ancestral trait for all of the extant allodapine lineages. Therefore comparisons among extant allodapines are unlikely to throw light on the initial origin of social behaviour, though they may uncover origins of true caste behaviour and reversals to solitary nesting. Sex ratios in Macrogalea and most other allodapine genera, spanning a broad phylogenetic and ecological range, suggest that one or more allodapine traits have provided persistent selection for female‐biased sex allocation.     

Colony Social Organisation and Alternative Social Strategies in the Eastern Carpenter Bee, <Emphasis Type="Italic">Xylocopa virginica</Emphasis>

The eastern carpenter bee, Xylocopa virginica, can nest either solitarily or in groups. In southern Ontario, Canada, near the northern edge of the range, most nests are social, containing groups of two to five adult females. Although social nests were much more frequent than solitary ones, they produced no more brood, so per capita brood productivity was actually lower for social females. Social females exhibited several reproductive strategies inferred from wing wear patterns, which reflect flight activity, and mandibular wear patterns, which reflect nest construction activity. Primary females accumulated a large degree of wing and mandibular wear and were presumed to be the primary reproductives in social nests. Secondary females accumulated less wing and mandibular wear and were probably subordinates awaiting opportunities to supersede primaries as dominant foragers and egg-layers. Tertiary females remained inactive, apparently deferring reproduction to the subsequent year. Social nesting, serial replacement of dominant females in social nests, and deferred reproduction are probably responses to severe competition for nests and nesting substrate.     

Strong constraints to independent nesting in a facultatively social bee: quantifying the effects of enemies-at-the-nest

Constraints to independent nesting play a key role in the understanding of social evolution in insects, but the source and the magnitude of such constraints are poorly known for many key taxa. In allodapine bees it is known that solitary nesting females have low rates of successful brood rearing and that this drives selection for cooperative nesting. It has been hypothesized that these constraints are due to the presence of enemies-at-the-nest, such as ants, but no direct link has been demonstrated between such enemies and colony failure. We set up an experiment in which solitary founded nests of an Australian allodapine bee, Exoneura nigrescens, were either protected from non-flying predators or left unprotected, and compared the resulting colony survival and brood production rates. We found that protected colonies have much higher rates of survival and that the constraints to independent nesting are extreme, with a mean of less than one offspring per nest at the end of the brood rearing period. This means that cooperative nesting is essential for this species to persist in its habitat. Received 6 July 2007; revised 5 November 2007; accepted 12 November 2007.     

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.     

Physiological variation as a mechanism for developmental caste-biasing in a facultatively eusocial sweat bee

Social castes of eusocial insects may have arisen through an evolutionary modification of an ancestral reproductive ground plan, such that some adults emerge from development physiologically primed to specialize on reproduction (queens) and others on maternal care expressed as allo-maternal behaviour (workers). This hypothesis predicts that variation in reproductive physiology should emerge from ontogeny and underlie division of labour. To test these predictions, we identified physiological links to division of labour in a facultatively eusocial sweat bee, Megalopta genalis. Queens are larger, have larger ovaries and have higher vitellogenin titres than workers. We then compared queens and workers with their solitary counterparts-solitary reproductive females and dispersing nest foundresses-to investigate physiological variation as a factor in caste evolution. Within dyads, body size and ovary development were the best predictors of behavioural class. Queens and dispersers are larger, with larger ovaries than their solitary counterparts. Finally, we raised bees in social isolation to investigate the influence of ontogeny on physiological variation. Body size and ovary development among isolated females were highly variable, and linked to differences in vitellogenin titres. As these are key physiological predictors of social caste, our results provide evidence for developmental caste-biasing in a facultatively eusocial bee.