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.     

Helper effects in the azure‐winged magpie Cyanopica cyana in relation to highly‐clumped nesting pattern and high frequency of conspecific nest‐raiding

In avian cooperative breeding systems, many benefits obtained by social pairs from the presence of helpers have been uncovered. However, until now, the factors that determine the type of assistance helpers provide and the responses of social pairs have not been well illustrated. We examined the contribution of helpers to cooperative groups and the relevant responses of dominant pairs in the azure‐winged magpie Cyanapica cyana which breeds on the Tibetan Plateau. We used the capture–mark–recapture method to identify helpers. Results showed that helpers were mostly the yearling sons of dominant pairs. They mainly contributed to the cooperative group in three ways, courtship‐feeding the incubation female, provisioning the brood, and defending the nest. For responses of dominant pairs, we unexpectedly found that clutch size was not influenced by the presence of helpers at the nest. However, cooperative groups had higher brood feeding rates than biparental nests and their feeding pattern also differed to that of the latter. Consequently, nestlings in cooperative groups had larger fledging body mass than that in biparental nests. By examining reasons for nest failure, we revealed that conspecific nest‐raiding contributed to more nest failure than any other natural predators. Because of the contribution of helpers in defending against both predators and conspecific nest‐raiders, cooperative groups had higher survival rate than biparental nests. Thus, our findings suggest that in a highly‐clumped nesting pattern, factors concerning the risk of nest predation, rather than that influencing food supply, play an important role in determining helper effects and responses of aided dominant pairs.     

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.     

Social Behaviour of the Carpenter Bee Xylocopa pubescens (Spinola)

The development of about 20 relatively small nests of Xylocopa pubescens was studied. After the first offspring had become adult, these nests reached a social stage in which there was only one egg-layer per nest. Freshly emerged (teneral) adults eat a lot of food, collected by a forager, before they fly out of the nest. This food appears to be of major importance to these bees in that it makes them fully agile. It seems therefore, that part of the food needed for the development of a young bee is not given at the larval stage, via the beebread, but at the teneral adult stage. As a result of this, a necessary overlap of generations is accomplished. Besides, less pollen has to be collected for the provision of a brood cell, so more cells can be constructed within a short period of time. There is a high degree of nest competition. Many nests were taken over by conspecific individuals (or by females of X. sulcatipes) that were searching for nesting sites within the study area. However, although more than 50% of the solitary nests were taken over sooner or later, strangers hardly ever intruded into social nests, with more than one adult. This illustrates how important it is for a reproducing female to tolerate the presence of nestmates which guard the nest in her absence. Although in most cases the foundress of a nest proceeded to produce new brood in the presence of her offspring, it happened that a daughter took over reproduction from her mother. In two of these cases, it could be observed that a mother started a new nest elsewhere after having been thrown out by her own daughter. At least in these cases, nest competition between mother and daughter started long before the mother had reached the end of her reproductive capacities. Since nesting possibilities are scarce, it seems a logical strategy to stay in the maternal nest and wait for a chance to become the egg-layer if the mother dies or if she loses her dominance. The occurrence of several social interactions among nestmates is discussed: — trophallaxis was observed often, not only under ‘forced’ conditions, but also as a form of ‘voluntary’ feeding; — nestmates were observed to groom each other.     

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).     

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.     

Colony Development and Serial Polygyny in the Primitively Eusocial Wasp Belonogaster juncea juncea (Vespidae: Polistinae)

In Cameroon, the social ropalidiine wasp Belonogaster juncea juncea mostly selects human constructions for nesting, but nests installed in the shelter of large boulders were also noted. Nests were founded throughout the year by one female (25.5%) or by associations of two to eight females. Among the 49 colonies studied from foundation, 24 produced at least one adult, and only 5 produced males among the eight that succeeded in producing sexual individuals. After the first dominant female disappeared (died or abandoned the nest, from 77 to 196 days after foundation) the remaining females fought among themselves, leading to a drastic decrease in the total number of brood. They completely abandoned 22 of 24 surviving nests, over a mean colony cycle lasting about 7 months. Hence the colony and nesting cycles were congruent, illustrating that determinate nesting corresponds to the majority of the cases in this species. In the other two nests, some females remained on the nest and began a new colonial cycle illustrating semi-independence between colony and nesting cycles. Up to four successive cycles were completed in this way in the same nest, thus illustrating the phenomenon of serial polygyny (intermediate between determinate and indeterminate nesting cycles). Serial polygyny in independent founding polistine wasps, previously documented only once, has implications for studies attempting to identify factors involved in regulating the colony and nesting cycles in tropical social wasps.     

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.     

Maternal investment in a bee species with facultative nest guarding and males heavier than females

1. Maternal investment can be influenced by several factors, especially maternal quality and possibilities for future reproduction. Mass provisioning Hymenoptera are an excellent group for measuring maternal investment because mothers distribute food sources to each brood cell for each offspring separately. Generally in aculeate Hymenoptera, larger females produce larger offspring and invest more in female offspring than in male offspring. 2. This study investigated patterns of maternal investment in Ceratina chalcites, which has an uncommon type of sexual size dimorphism in Hymenoptera: on average, males are heavier than females. It was found that larger females produce a significantly higher proportion of male offspring, as males are the costlier sex in this species. 3. Facultative nest guarding by females was observed. Females can guard offspring until adulthood, as is typical for bees of genus Ceratina (34.43% of nests); however, in the majority of cases (65.56% of nests), females plug and abandon the nest. Significant differences were found in the amount of investment between guarded and unguarded nests. Guarded nests had a greater number of provisioned brood cells and a higher proportion of male offspring. It is suggested that mothers have two facultative strategies – either she makes a large investment in the offspring of one nest or she abandons the first nest and carries out a second nesting elsewhere.     

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.     

Social polymorphism in the Australian small carpenter bee, Ceratina (Neoceratina) australensis

The bee tribe Ceratinini provides important insights into the early stages of sociality. Many arthropods exhibit behaviours precursory to social life, including adult longevity, parental care, nest loyalty and mutual tolerance, yet the origins of social behaviour are few. Here we describe the biology of a facultatively social bee, Ceratina (Neoceratina) australensis, which exhibits all of the preadaptations for successful group living, yet presents ecological and behavioural characteristics that seemingly disfavour frequent colony formation. This species is socially polymorphic with both solitary and social nests collected in sympatry. Social colonies consist of two foundresses, one contributing both foraging and reproductive effort and the second which remains at the nest as a passive guard. Cooperative nesting provides no overt reproductive benefits over solitary nesting in this population, although brood survival tends to be greater in social colonies. Maternal longevity, subsociality and bivoltine nesting phenology in this species favour colony formation, while dispersal habits and offspring longevity may inhibit more frequent social nesting in this and other ceratinines.     

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.     

Interactions between the Social Digger Wasp, <Emphasis Type="Italic">Cerceris rubida</Emphasis>, and Its Brood Parasitic Flies at a Mediterranean Nest Aggregation

The fitness consequence of many behaviours of the small digger wasp, Cerceris rubida (Hymenoptera: Crabronidae), the only European species of its genus in which females share nests, are still unknown. Here, I present novel data on the nesting patterns and nest parasites of an Italian population of this wasp, with emphasis on which behavioural strategies may have evolved to reduce brood parasitism. Nests were established mainly in horizontal surfaces with scarce vegetation and hard soil, resulting in spatially clumped nests; the extent of nest aggregation increased over a 6-year period. Wasp brood cells are attacked by the miltogrammine fly Pterella grisea (Diptera: Sarcophagidae), which waits for nest-returning wasps on perching sites and then follows them in flight (satellite flight), eventually landing on the nest entrance and larvipositing without entering further in the tunnel. This technique seems to be adaptive for the parasitic flies, which would be rejected from nests by the guarding wasps if attempting to enter. The daily activity of the flies closely matched the host wasp’s provisioning activity, but C. rubida females were able to partially confound the tracking flies by performing evasive manoeuvres while returning to the nest. Patches with higher nest density and nests with more resident females did suffer more fly landings on nest entrances (a prerequisite for larviposition). These trends, however, disappeared on a per nest basis and on a per wasp provisioning flight basis, respectively. Across two years, only 6% of brood cells were parasitized, a picture similar to what observed for miltogrammine flies at nest aggregations of other Cerceris spp., and new data are necessary to test if there is a benefit of either nest density or nest sharing against P. grisea parasitism.     

Effects of nest-site characteristics and parental activity on cowbird parasitism and nest predation in Brown-and-yellow Marshbirds

ABSTRACT Nest‐site selection and nest defense are strategies for reducing the costs of brood parasitism and nest predation, two selective forces that can influence avian nesting success and fitness. During 2001–2002, we analyzed the effect of nest‐site characteristics, nesting pattern, and parental activity on nest predation and brood parasitism by cowbirds (Molothrus spp.) in a population of Brown‐and‐yellow Marshbirds (Pseudoleistes virescens) in the Buenos Aires province, Argentina. We examined the possible effects of nest detectability, nest accessibility, and nest defense on rates of parasitism and nest predation. We also compared rates of parasitism and nest predation and nest survival time of marshbird nests during the egg stage (active nests) with those of the same nests artificially baited with passerine eggs after young fledged or nests failed (experimental nests). Most nests (45 of 48, or 94%) found during the building or laying stages were parasitized, and 79% suffered at least one egg‐predation event. Cowbirds were responsible for most egg predation, with 82 of 107 (77%) egg‐predation events corresponding to eggs punctured by cowbirds. Nests built in thistles had higher rates of parasitism and egg predation than nests in other plant, probably because cowbirds were most active in the area where thistles were almost the only available nesting substrate. Parasitism rates also tended to increase as the distance to conspecific nests increased, possibly due to cooperative mobbing and parental defense by marshbirds. The proportion of nests discovered by cowbirds was higher for active (95%) than for experimental (29%) nests, suggesting that cowbirds used host parental activity to locate nests. Despite active nest defense, parental activity did not affect either predation rates or nest‐survival time. Thus, although nest defense by Brown‐and‐yellow Marshbirds appears to be based on cooperative group defense, such behavior did not reduce the impact of brood parasites and predators.     

Sulcopolistes atrimandibularis,Social Parasite and Predator of an Alpine Polistes (Hymenoptera,Vespidae)1

Sulcopolistes atrimandibularis Zimmermann is the obligate social parasite of Polistes biglumis bimaculatus Geoffry in Furcroy, a mountain species of paper wasp. Unlike all the other hymenopteran social parasites, the Sulcopolistes female obtains part of the food for her immature brood by plundering other nests of the host species. Parasite females can control more than one host nest: one of them (the nursery nest) she uses solely for reproduction purposes (from which the Sulcopolistes reproductives emerge) and others (supply nests) are used for exploiting the Polistes brood content. It is possible that this behaviour is an adaptation to the extremely short colonial cycle of the host.     

Host-parasite relationships in six species ofSphecodes bees and their halictid hosts: Nest intrusion,intranidal behavior,and Dufour's gland volatiles (Hymenoptera: Halictidae)

Nest invasion behavior was studied in six kleptoparasiticSphecodes species at four nesting sites of their respective social and solitary hosts.Sphecodes females preferred to enter unguarded nests. Nest intruding strategies observed in the differentSphecodes species did not depend on whether host species were solitary or social, as long as the nesting cycle of a social host was in the solitary stage (i.e., a single host female). Observation of intranidal behavior revealed thatSphecodes monilicornis females kill all host individuals within an usurped nest. They stay in the nest for several hours, laying eggs in adequately provisioned brood cells. Gas chromatography-mass spectrometry analyses of Dufour's gland secretions revealed species-specific compositions. Qualitative comparisons of whole patterns and quantitative comparisons considering the predominant hydrocarbons common to both host and parasite contradict the hypothesis of chemical mimetism, a mechanism supposed to permit parasite intrusion by qualitatively similar odor bouquets in host and parasite females.     

Kin grouping is insufficient to explain the inclusive fitness gains of conspecific brood parasitism in the common eider

Conspecific brood parasitism allows females to exploit other females' nests and enhance their reproductive output. Here, we test a recent theoretical model of how host females gain inclusive fitness from brood parasitism. High levels of relatedness between host and parasitizer can be maintained either by: (a) kin recognizing and parasitizing each other as a form of cooperative breeding or (b) natal philopatry and nest site fidelity facilitating the formation of kin groups, thereby increasing the probability of parasitism between relatives nesting in close proximity. To address these two hypotheses we genotyped feathers and hatch membranes of common eiders (Somateria mollissima) from western Hudson Bay, Canada, using a noninvasive sampling methodology. We found that most instances of brood parasitism do result in inclusive fitness gains. Furthermore, females with failed nests moved an average of 492 m from their previous year's nest site, while successful females only moved an average of 13 m. Therefore, we observed host–parasite relatedness can occur at levels higher than would be expected by chance even in the absence of kin grouping, suggesting that closely related females nesting near one another is not essential to maintain high host–parasitizer relatedness. In addition, kin grouping is only a transient phenomenon that cannot occur every year due to the propensity for females of failed nests to nest farther away from their nest site in subsequent years than females with successful nests, which provides support for kin recognition as a more likely mechanism to maintain high host–parasitizer relatedness over time.     

FEMALE-BIASED SEX RATIOS IN A FACULTATIVELY SOCIAL BEE AND THEIR IMPLICATIONS FOR SOCIAL EVOLUTION

Montane populations of the Australian allodapine bee, Exoneura bicolor, are characterized by high levels of cooperative nesting and strongly female-biased sex ratios. A conspecific population from heathland shows much lower levels of cooperative nesting and lower levels of female bias. In both habitats, sex-ratio bias is greatest in the smallest brood sizes and becomes successively less biased in larger broods. Parity is approached in the largest heathland colonies, but not for any brood-size category in montane areas. Adult intracolony relatedness is moderately high for colonies in both reused and newly founded nests in the montane habitat, but probably low or zero for newly founded nests in heathland. Colony efficiency, measured as the number of brood per adult, increases with colony size in both habitats, suggesting that cooperation between females increases mean female fitness. It is argued that patterns of sex allocation are consistent with nonlinear fitness-return models, in which the mean reproductive value of daughters increases with the number of daughters produced in a brood. Such increases probably arise from a number of social interactions, including cooperative brood defense, increased task efficiency, and lower per capita costs in nest construction. The term “local fitness enhancement” is introduced here to describe these effects collectively. The female-biased ratios should lower selective thresholds for sib-directed altruism, at least in the earlier stages of colony development. It is argued that local fitness enhancement facilitates eusociality in allodapine bees and could also play a role in other haplodiploid taxa, provided cooperative nesting largely involves sisters, colony efficiency increases with colony size, and optimal colony sizes are only achieved after two or more generations after founding.     

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.