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Abstract Insect parental care strategies are particularly diverse, and prolonged association between parents and offspring may be a key precursor to the evolution of complex social traits. Macroevolutionary patterns remain obscure, however, due to the few rigorous phylogenetic analyses. The subsocial sphecid wasps are a useful group in which to study parental care because of the diverse range of strategies they exhibit. These strategies range from placing a single prey item in a pre-existing cavity to mass provisioning a pre-built nest, through to complex progressive provisioning where a female feeds larvae in different nests simultaneously as they grow. We show that this diversity stems from multiple independent transitions between states. The strategies we focus on were previously thought of in terms of a stepping-stone model in which complexity increases during evolution, ending with progressive provisioning which is a likely precursor to eusociality. We find that evolution has not always followed this model: reverse transitions are common, and the ancestral state is the most flexible rather than the simplest strategy. Progressive provisioning has evolved several times independently, but transitions away from it appear rare. We discuss the possibility that ancestral plasticity has played a role in the evolution of extended parental care. Significance statement Parental care behaviour leads to prolonged associations between parents and offspring, which is thought to drive the evolution of social living. Despite the importance of insect parental care for shaping the evolution of sociality, relatively few studies have attempted to reconstruct how different strategies evolve in the insects. In this study, we use phylogenetic methods to reconstruct the evolution of the diverse parental care strategies exhibited by the subsocial digger wasps (Sphecidae). Contrary to expectations, we show that parental care in this group has not increased in complexity over evolutionary time. We find that the ancestral state is not the simplest, but may be the most flexible strategy. We suggest that this flexible ancestral strategy may have allowed rapid response to changing environmental conditions which might explain the diversity in parental care strategies that we see in the digger wasps today.

Understanding the ecological and environmental contexts in which eusociality can evolve is fundamental to elucidating its evolutionary origins. A sufficiently long active season is postulated to have been a key factor facilitating the transition to eusociality. Many primitively eusocial species exhibit an annual life cycle, which is thought to preclude the expression of eusociality where the active season is too short to produce successive worker and reproductive broods. However, few studies have attempted to test this idea experimentally. We investigated environmental constraints on the expression of eusociality in the obligate primitively eusocial sweat bee Lasioglossum malachurum, by transplanting nest foundresses from the south to the far north of the United Kingdom, far beyond the natural range of L. malachurum. We show that transplanted bees can exhibit eusociality, but that the short length of the season and harsher environmental conditions could preclude its successful expression. In one year, when foundresses were transplanted only after provisioning first brood (B1) offspring, workers emerged in the north and provisioned a second brood (B2) of reproductives. In another year, when foundresses were transplanted prior to B1 being provisioned, they were just as likely to initiate nesting and provisioned just as many B1 cells as foundresses in the south. However, the life cycle was delayed by approximately 7 weeks and nests suffered 100% B1 mortality. Our results suggest that short season length together with poor weather conditions represent an environmental barrier to the evolution and expression of eusociality in sweat bees. Electronic supplementary material The online version of this article (10.1007/s00040-018-0642-7) contains supplementary material, which is available to authorized users.

Cooperation among kin is common in animal societies. Kin groups may form by individuals directly discriminating relatives based on kin recognition cues, or form passively through natal philopatry and limited dispersal. We describe the genetic landscape for a primitively eusocial wasp, Polistes dominula, and ask whether individuals choose cooperative partners that are nearby and/or that are genetic relatives. Firstly, we genotyped an entire sub-population of 1361 wasps and found genetic structuring on an extremely fine scale: the probability of finding genetic relatives decreases exponentially within just a few meters of an individual’s nest. At the same time, however, we found a lack of genetic structuring between natural nest aggregations within the population. Secondly, in a separate dataset where ~2000 wasps were genotyped, we show that wasps forced experimentally to make a new nest choice tended to choose new nests near to their original nests, and that these nests tended to contain some full sisters. However, a significant fraction of wasps chose nests that did not contain sisters, despite sisters being present in nearby nests. Although we cannot rule out a role for direct kin recognition or natal nest-mate recognition, our data suggest that kin groups may form via a philopatric rule-of-thumb, whereby wasps simply select groups and nesting sites that are nearby. The result is that most subordinate helpers obtain indirect fitness benefits by breeding cooperatively.

In cooperative breeders, subordinates that have alternative reproductive options are expected to stay and help dominant breeders only as long as they contribute to group productivity, if their fitness is linked with colony success. Female Polistes dominula paper wasps live as cooperative breeders in small groups of typically fewer than 10 females. Subordinates tend to have high-quality outside options, and so could choose alternative breeding tactics if their work efforts increased productivity negligibly. In the founding stage before workers emerge, we tested the effect of various predictors on nest growth, as a proxy for group productivity, and explored the shape of the relationship between group size and nest growth. We found group size to be the only significant predictor of nest growth: variation among body sizes within the group showed no effect, suggesting a lack of size-dependent task specialization in this species. Average body size and average genetic relatedness between group members similarly showed no effects on nest growth. Group size had a non-linear effect so that per-capita benefits to nest growth decreased in larger groups, and groups of 10 or more would benefit negligibly from additional group members. Hence, females might be better off pursuing other options than joining a large group. This finding helps to explain why P. dominula groups are usually relatively small in our study population. Further studies may illuminate the mechanisms behind the smaller per-capita nest growth that we found in larger groups. Significance statement Identifying which factors influence the productivity of animal groups is key to understanding why different species breed cooperatively in groups of varying sizes. In the paper wasp Polistes dominula, we investigated the growth rate of nests as a measure of group productivity. We found that average body size, the variation in body sizes within the group, and average genetic relatedness between group members did not affect nest growth, while group size had a strong, positive effect: nests grew faster with more group members, but the per-capita benefit decreased in larger groups. The addition of extra group members in groups of 10 or more had negligible effects on nest growth. Hence, wasps may be better off pursuing other options than joining large groups. This finding helps to explain why groups normally consist of fewer than 10 wasps in this population. Electronic supplementary material The online version of this article (10.1007/s00265-018-2502-x) contains supplementary material, which is available to authorized users.

Eusociality is characterised by a reproductive division of labour, where some individuals forgo direct reproduction to instead help raise kin. Socially polymorphic sweat bees are ideal models for addressing the mechanisms underlying the transition from solitary living to eusociality, because different individuals in the same species can express either eusocial or solitary behaviour. A key question is whether alternative social phenotypes represent environmentally induced plasticity or predominantly genetic differentiation between populations. In this paper, we focus on the sweat bee Lasioglossum calceatum, in which northern or high-altitude populations are solitary, whereas more southern or low-altitude populations are typically eusocial. To test whether social phenotype responds to local environmental cues, we transplanted adult females from a solitary, northern population, to a southern site where native bees are typically eusocial. Nearly all native nests were eusocial, with foundresses producing small first brood (B1) females that became workers. In contrast, nine out of ten nests initiated by transplanted bees were solitary, producing female offspring that were the same size as the foundress and entered directly into hibernation. Only one of these ten nests became eusocial. Social phenotype was unlikely to be related to temperature experienced by nest foundresses when provisioning B1 offspring, or by B1 emergence time, both previously implicated in social plasticity seen in two other socially polymorphic sweat bees. Our results suggest that social polymorphism in L. calceatum predominantly reflects genetic differentiation between populations, and that plasticity is in the process of being lost by bees in northern populations. Phenotypic plasticity is thought to play a key role in the early stages of the transition from solitary to eusocial behaviour, but may then be lost if environmental conditions become less variable. Socially polymorphic sweat bees exhibit either solitary or eusocial behaviour in different geographic populations, depending on the length of the nesting season. We tested for plasticity in the socially polymorphic sweat bee Lasioglossum calceatum by transplanting nest foundresses from a northern, non-eusocial population to a southern, eusocial population. Plasticity would be detected if transplanted bees exhibited eusocial behaviour. We found that while native bees were eusocial, 90% of transplanted bees and their offspring did not exhibit traits associated with eusociality. Environmental variables such as time of offspring emergence or temperatures experienced by foundresses during provisioning could not explain these differences. Our results suggest that the ability of transplanted bees to express eusociality is being lost, and that social polymorphism predominantly reflects genetic differences between populations.

Recent theory and empirical studies of avian biparental systems suggest that animals resolve conflict over parental care via a process of behavioral negotiation or “rules for responding.” Less is known, however, about whether negotiation over helping effort occurs in cooperatively breeding animal societies or whether behavioral negotiation requires a relatively large brain. In this study, we tested whether negotiation over help occurs in a social insect, the paper wasp Polistes dominulus, by recording individual responses to both observed and experimentally induced foraging returns by other group members. In our experiments, we manipulated food delivery to the nest in 2 ways: 1) by catching departing foragers and giving them larval food to take back to the nest and 2) by giving larval food directly to wasps on the nest, which they then fed to larvae, so increasing food delivery independently of helper effort. We found no evidence from Experiment 1 that helpers adjusted their own foraging effort according to the foraging effort of other group members. However, when food was provided directly to the nest, wasps did respond by reducing their own foraging effort. One interpretation of this result is that paper wasp helpers adjust their helping effort according to the level of offspring need rather than the work rate of other helpers. Negotiation based on indicators of demand rather than work rate is a likely mechanism to resolve conflict over investment in teams where helpers cannot observe each other’s work rate directly, as is commonly the case in insect and vertebrate societies.

Relatedness is predicted to be a key determinant of cooperative behavior, but kin discrimination within social insect colonies is surprisingly rare. A lack of reliable cuticular hydrocarbon (CHC) cues is thought to be responsible, but here we show that in a high-profile paper wasp model, kin recognition cues are available for some individuals that found nests with nonrelatives. Thus, unrelated Polistes dominulus helpers could potentially recognize themselves as such. On this basis, we reanalyzed a behavioral data set to investigate whether foraging effort, defense contributions and aggression toward nest mates might thus reflect CHC profiles. Both foraging behavior and aggression varied with genetic relatedness, but genetic relatedness itself was a better predictor of this variation than differences in CHC profiles. We propose that wasps use specific components of the CHC profile, the identity of which is as yet unknown, to identify relatives among nest mates. Our data provide the first evidence of within-nest kin discrimination in primitively eusocial wasps but leave open the question of which cues are responsible.

Within-group power asymmetries and the resulting reproductive skew, common in most social groups, may effectively be set at the very early stages of group formation, that is, when group membership is determined. Hence, groupmate choices can define an individual’s future reproductive success. We examined how groups of Polistes dominulus formed under natural, unconstrained conditions, using data on the nesting history, kinship and morphology of individually marked foundresses obtained during two consecutive seasons in southern Spain. Foundresses that hibernated in the same aggregation were more likely to start a nest together, but all of the foundresses at a nest were seldom from a single aggregation. Changes in group composition were frequent throughout the preworker period, mainly because some foundresses disappeared and other wasps joined established groups. Within-group relatedness, however, was not affected by the late arrival of wasps. Our results suggest that waiting to join an established group is a common nesting strategy in P. dominulu s. Only 16% of marked wasps used more than one nest. Foundresses that moved between groups tended to move to groups in which genetic relatedness among the resident foundresses was higher, but not necessarily relatedness to the moving wasp herself. Overall, nestmate choices were not associated with a single factor. High failure rates, particularly of single-foundress nests, however, suggest that ecological constraints (e.g. risk of predation, lack of resources) may have a stronger effect on individual nesting choices than previously considered.

Altruism in the social Hymenoptera is generally considered to be a feature of females rather than males. A popular explanation for this is that in the solitary ancestors of today's social species, males provided little brood care. Males might therefore lack the preadaptations necessary to evolve altruism in social contexts. While anecdotal observations of male contributions to colony life have been reported, there are few reports of male participation in nest defence. In apoid wasps, there have been several reports of male nest-guarding behaviour in solitary species, potentially setting the evolutionary stage for similar behaviours in social lineages. Here, we present evidence of active and effective nest defence in males of the social apoid wasp Microstigmus nigrophthalmus. Males were observed chasing intruders away from the nest, and the presence of males had a significant effect on nest survival when females were removed. Males potentially obtained direct benefits through defence, so that defence may not represent male altruism. However, our results do show that males can perform acts that benefit their colony.

Our knowledge of primitively eusocial societies is focused particularly on cooperatively breeding vertebrates and vespid wasps, whereas numerous taxa representing independent origins of social behavior have been largely overlooked. The lineage of apoid wasps including the genus Microstigmus represents a relatively neglected independent origin of eusociality. We present the first use of modern hypervariable genetic markers, in combination with behavioral observations, to investigate reproductive division of labor and cooperative brood care in an apoid wasp, the Brazilian M. nigrophthalmus. Microstigmus nigrophthalmus is unusual because, although there is cooperative brood care, reproductively dominant females carry out at least as much risky foraging as their subordinate nest mates. Empirical studies of reproductive skew are often hampered by a lack of variation in skew. We find that reproductive skew is highly variable between nests in M. nigrophthalmus. There was no correlation between skew and either body size or group size. The absence of an effect of body size is typical of studies of skew in insects and may indicate that body size is a poor measure of an individual's ability to control a group. However, skew was positively correlated with genetic relatedness. This provides rare support for concession models based on "social contracts" between dominant individuals and their subordinates. Copyright 2011, Oxford University Press.

Environmental factors, and particularly climate, play an important role in influencing behaviour in many insects. In social species, climate is known to have a strong influence on social traits, but how this manifests itself in tropical ecosystems is poorly understood. In Peninsular Malaysia, the climate is characterised by relatively consistent annual temperatures with wet/dry cycles, and the tropical hover-wasp Liostenogaster flavolineata Cameron is active year-round. Newly emerged females can choose to remain at the natal nest and help, or disperse and found their own nest depending on a balance of ecological and demographic factors. We collated long-term adult and brood census data for populations of L. flavolineata in Peninsular Malaysia in three different years to investigate temporal variation in demographics (brood and adult numbers) and how this might be related to climatic factors. Our data indicate that there are multiple, temporally distinct peaks of brood production in this population. The number of newly eclosing females and number of mature brood were positively associated with temperature and negatively associated with the number of rain-days during the observation period. Furthermore, larger females were produced during the peaks of brood production. We speculate how these patterns may influence the staying or leaving decisions of newly emerged females in a primitively eusocial species such as L. flavolineata.

Hierarchies constitute the base of many social groups. Hence, understanding how they are established is critical. Here we examine how hierarchies are formed in foundresses associations of the common paper wasp Polistes dominulus. By comparing field data with computer simulations, we evaluate order of arrival at the nest, body size, facial color patterns, and within-group kinship structure as determinants of inheritance rank. Hierarchies (ranks 1--5) were experimentally inferred for 53 nests. Overall, the order in which foundresses arrived at the nest and their body size were not significantly correlated with rank. A foundress's rank was negatively correlated with the number of full sisters it had in its group. Highly ranked wasps (ranks 1 and 2) were less likely to share a nest with their full sisters than wasps of lower rank. A wasp's rank was not determined by the relative rank of its nest-mate sisters. A foundress's rank was significantly correlated with the size of its black clypeal marks, but the number of foundresses with clypeal marks in each nest was small. On 15 of 20 nests where wasps with marks were present, only 1 wasp had such marks. Overall, our results suggest that within-group relatedness structure is important in the establishment of dominance hierarchies in P. dominulus foundress associations. Copyright 2009, Oxford University Press.

The sweat bees (Family Halictidae) are a socially diverse taxon in which eusociality has arisen independently numerous times. The obligate, primitively eusocial Lasioglossum malachurum, distributed widely throughout Europe, has been considered the zenith of sociality within halictids. A single queen heads a colony of smaller daughter workers which, by mid-summer, produce new sexuals (males and gynes), of which only the mated gynes overwinter to found new colonies the following spring. We excavated successfully 18 nests during the worker- and gyne-producing phases of the colony cycle and analysed each nest's queen and either all workers or all gynes using highly variable microsatellite loci developed specifically for this species. Three important points arise from our analyses. First, queens are facultatively polyandrous (queen effective mating frequency: range 1?3, harmonic mean 1.13). Second, queens may head colonies containing unrelated individuals (n= 6 of 18 nests), most probably a consequence of colony usurpation during the early phase of the colony cycle before worker emergence. Third, nonqueen's workers may, but the queen's own workers do not, lay fertilized eggs in the presence of the queen that successfully develop into gynes, in agreement with so-called `concession? models of reproductive skew.

The mechanisms through which dominance is inherited within social groups vary from direct interactions such as fighting to non-confrontational conventions. Liostenogaster flavolineata is a primitively eusocial hover wasp in which one female, the ‘dominant’, is the only reproductive upon the nest. The remaining females, although capable of reproduction, behave as helpers. In this study, we investigate the rules by which helpers inherit dominance. We removed successive dominants from 56 nests and recorded accession on un-manipulated nests. The results showed that L. flavolineata has a strict age-based inheritance queue: new dominants are the oldest female in their groups 87% of the time. Thirteen cases of queue-jumping were found in which young individuals were able to supplant older nestmates and inherit dominance precociously. Queue jumpers did not differ from other wasps in terms of relatedness to other group members or body size. Individuals that had previously worked less hard than other females of equivalent rank were significantly more likely to later jump the queue. Queue-jumping may represent a cheating strategy or could indicate that the rule for inheriting dominance is not based purely on relative age. We also discuss possible reasons why age-based queuing has evolved and its potential to promote the evolution of helping behaviour.

An important benefit of social living is increased capacity for defense. Highly eusocial species have often evolved a fighting caste for this purpose, but many facultatively eusocial insects and cooperatively breeding vertebrates lack morphological castes and the decision to defend or not can depend on costs and benefits to each individual. Defense by subordinates in a social group can be regarded as a form of helping, and helping input often varies among subordinates of different age or size. Several hypotheses attempt to explain variation in helping effort, including the effects of relatedness and differences in the costs of helping. Evidence for these hypotheses is mixed and often lacks data on the rank of subordinates, an important determinant of expected future fitness. We examined individual variation in propensity to defend the nest against conspecifics in the tropical hairy-faced hover wasp Liostenogaster flavolineata. Prior to experimentation, we determined the positions of all wasps in the age-based queue to inherit the single egg-laying position in each L. flavolineata group. Two approaches were then used: observations of defense against natural intrusions by conspecifics and experimental trials where wasps were presented attached to a wire. Higher ranks were more likely to defend the nest than lower ranks, opposite to the pattern previously documented for another form of helping: foraging effort. Possible explanations for this result are that higher ranked females are better defenders and that they suffer a larger decrease in expected future fitness when an intruder usurps their position in the inheritance queue. Copyright 2007, Oxford University Press.

Status signals function in a number of species to communicate competitive ability to conspecific rivals during competition for resources. In the paper wasp Polistes dominulus, variable black clypeal patterns are thought to be important in mediating competition among females. Results of previous behavioral experiments in the lab indicate that P. dominulus clypeal patterns provide information about an individual’s competitive ability to rivals during agonistic interactions. To date, however, there has been no detailed examination of the adaptive value of clypeal patterns in the wild. To address this, we looked for correlations between clypeal patterning and various fitness measures, including reproductive success, hierarchical rank, and survival, in a large, free-living population of P. dominulus in southern Spain. Reproductive success over the nesting season was not correlated with clypeal patterning. Furthermore, there was no relationship between a female’s clypeal patterning and the rank she achieved within the hierarchy or her survival during nest founding. Overall, we found no evidence that P. dominulus clypeal patterns are related to competitive ability or other aspects of quality in our population. This result is consistent with geographical variation in the adaptive value of clypeal patterns between P. dominulus populations; however, data on the relationship between patterning and fitness from other populations are required to test this hypothesis. Key wor ds: clypeal pattern, fitness, Polistes dominulus, quality, status signalling, wild. [Behav Ecol]

The functional significance of nest co-occupation inCerceris arenaria(Hymenoptera: Sphecidae) was investigated. Willmer (1985a,Behav. Ecol. Sociobiol.,17,151-160, 1987,Proc. 3rd Eur. Congr. Entomol,2,229-240) suggested that in this species co-occupation is a mutualistic response to ecophysiological pressures, and that these pressures might have played a general role in the evolution of eusociality. In this study nest-less ‘Searcher’ females almost always (99%,N=80) accepted (started using) pre-existing nests rather than digging new burrows. Burrows were occupied by sequences of females whose periods of occupation overlapped briefly. Co-occupations began when Searchers accepted nests that already had owners. The behaviour of Searchers after nest acceptance depended on time of acceptance. Late in the day, Searchers remained inside nests and usually closed them. Early in the day, Searchers started provisioning accepted nests. If these nests were already occupied, the result was joint-provisioning. When a nest was joint-provisioned, it received significantly more prey than the owner could provision alone, but less than twice what she could provision alone. Co-occupations rarely (4%,N=47) lasted more than 1 day, and ended when one female left the nest for good. Searchers remained in possession of nests after 49% (N=47) of co-occupations. Searchers appear functionally to be usurpers: they are known to prefer nests with owners and on average, owners lose out if Searchers accept their nests. Willmer's hypothesis that co-occupations are mutualistic is rejected. An analysis of time costs suggests that accepting a pre-existing nest has larger payoffs than digging a new burrow. The high rate of replacement of nest owners (11-16% per day) will reduce the relatedness of offspring emerging from the same nest, presenting a barrier against the kin-selected evolution of nest-sharing. Variation in social interactions in the genusCercerismay be consistent with the predictions of a recent model of cooperation based on reciprocity.

Climate has long been suggested to affect population genetic structures of eusocial insect societies. For instance, Hamilton [Journal of Theoretical Biology 7 (1964) 17] discusses whether temperate and tropical eusocial insects may show differences in population-level genetic structure and viscosity, and how this might relate to differences in the degree of synchrony in their life cycles or modes of nest founding. Despite the importance of Hamilton's 1964 papers, this specific idea has not been tested in actual populations of wasps, probably due to the paucity of studies on tropical species. Here, we compare colony and population genetic structures in two species of primitively eusocial paper wasps with contrasting ecologies: the tropical species Polistes canadensis and the temperate species P. dominulus. Our results provide important clarifications of Hamilton's discussion. Specifically, we show that the genetic structures of the temperate and tropical species were very similar, indicating that seasonality does not greatly affect population viscosity or inbreeding. For both species, the high genetic differentiation between nests suggests strong selection at the nest level to live with relatives, whereas low population viscosity and low genetic differentiation between nest aggregations might reflect balancing selection to disperse, avoiding competition with relatives. Overall, our study suggests no prevalence of seasonal constraints of the life cycle in affecting the population genetic structure of eusocial paper wasps. These conclusions are likely to apply also to other primitively eusocial insects, such as halictine bees. They also highlight how selection for a kin structure that promotes altruism can override potential effects of ecology in eusocial insects. © 2012 The Authors.Ecology and Evolution published by Blackwell Publishing Ltd.

Abstract. This paper demonstrates that intraspecific nest parasites are selective in their choice of host nests. In a large nest aggregation, instead of digging new burrows, nest-less females of the digger wasp Cerceris arenaria entered pre-existing burrows and eventually used one of them. Data on host nest selection showed that the likelihood that a nest-less female would use a burrow increased if the burrow contained more prey, if the burrow had a conspecific owner, and if the owner was absent from the nest. This indicates that during their brief visits to burrows, nest-less females are capable of rapidly assessing the value of the burrow and its contents. They appear to be true 'usurpers', unlike nest-less females of other species which use burrows at random ('entering'). The evolution of an ability to assess prey may not have involved large costs since a similar ability may have already been present, associated with provisioning and sex allocation strategies. A significant cost of intraspecific parasitism could arise if the same nests tend to be repeatedly parasitized, but this was not the case in C. arenaria.

Animals that cooperate with nonrelatives represent a challenge to inclusive fitness theory, unless cooperative behavior is shown to provide direct fitness benefits. Inheritance of breeding resources could provide such benefits, but this route to cooperation has been little investigated in the social insects. We show that nest inheritance can explain the presence of unrelated helpers in a classic social insect model, the primitively eusocial wasp Polistes dominulus. We found that subordinate helpers produced more direct offspring than lone breeders, some while still subordinate but most after inheriting the dominant position. Thus, while indirect fitness obtained through helping relatives has been the dominant paradigm for understanding eusociality in insects, direct fitness is vital to explain cooperation in P. dominulus.

The theory of assured fitness returns proposes that individuals nesting in groups gain fitness benefits from effort expended in brood-rearing, even if they die before the young that they have raised reach independence. These benefits, however, require that surviving nest-mates take up the task of rearing these young. It has been suggested that assured fitness returns could have favoured group nesting even at the origin of sociality (that is, in species without a dedicated worker caste). We show that experimentally orphaned brood of the apoid wasp Microstigmus nigrophthalmus continue to be provisioned by surviving adults for at least two weeks after the orphaning. This was the case for brood of both sexes. There was no evidence that naturally orphaned offspring received less food than those that still had mothers in the nest. Assured fitness returns can therefore represent a real benefit to nesting in groups, even in species without a dedicated worker caste.

The paper wasp Polistes dominulus is unique among the social insects in that nearly one-third of co-foundresses are completely unrelated to the dominant individual whose offspring they help to rear and yet reproductive skew is high. These unrelated subordinates stand to gain direct fitness through nest inheritance, raising the question of whether their behaviour is adaptively tailored towards maximizing inheritance prospects. Unusually, in this species, a wealth of theory and empirical data allows us to predict how unrelated subordinates should behave. Based on these predictions, here we compare helping in subordinates that are unrelated or related to the dominant wasp across an extensive range of field-based behavioural contexts. We find no differences in foraging effort, defense behaviour, aggression or inheritance rank between unrelated helpers and their related counterparts. Our study provides no evidence, across a number of behavioural scenarios, that the behaviour of unrelated subordinates is adaptively modified to promote direct fitness interests.\ud \ud

The nesting behaviour of individually marked female pompilid wasps, Anoplius viaticus, was observed at a Breckland heath site with particular emphasis on intraspecific parasitism and nest defence. Prey was stolen from conspecifics while it was being carried to the nest site, and while it was left unattended during nest construction. Females also appeared to brood parasitize each other's completed nests. Parasitism appeared to be opportunistic. Brood parasitism may be a tactic by which time-limited females can increase their fecundity. By placing prey in vegetation tufts during nest construction, females may reduce the risk of prey theft. An individual female's successive unicellular nests were clustered and therefore easier to defend, in many ways resembling a multicellular nest. Females defended their clusters vigorously, visiting them every few minutes during foraging and expelling conspecifics from the vicinity. This type of nest defence may be costly, and has rarely been observed in solitary wasps.

1 In nest-building wasps there is a continuum from species that provision each offspring cell with one large prey item (single provisioning) to species that provision each cell with many small items (multiple provisioning). The significance of number of prey per cell was examined during two seasons in Ammophila sabulosa (L.), a species that provisions between one and five caterpillars per cell. 2 There was no difference between the total weight of prey in singly-provisioned and multiply-provisioned cells: individual caterpillars placed in multiply-provisioned cells were smaller. 3 Small caterpillars were captured and transported to the nest faster than large ones. 4 If the first prey provisioned was large, the cell was usually then permanently closed. If the first prey was small, additional prey were captured, and females appeared to become more selective: additional prey were rarely large. 5 It is suggested that the flexible provisioning strategy used by A.sabulosa allows a wide range of prey sizes to be utilized without affecting maternal control of offspring size and sex. This could be important when available prey sizes vary temporally and spatially. 6 There was a male-biased first generation investment ratio. Female-producing cells were provisioned with a higher total prey weight than male-producing cells, and conversion of prey weight to adult weight was more efficient for females. 7 Total prey weight provisioned is probably a good indirect measure of parental time investment in A.sabulosa. Other measures, particularly number of prey provisioned, will be less accurate.

Nest-building Hymenoptera have been a major testing ground for theories of parental investment and sex allocation. Investment has usually been estimated by the likely costs of offspring provisioning, ignoring other aspects of parental care. Using three experimental treatments, we estimated the costs of egg-laying and provisioning separately under field conditions in a digger wasp Ammophila pubescens . In one treatment, we increased the provisioning effort required per offspring by removing alternate prey items as they were brought to the nest. In two other treatments, we reduced parental effort by either preventing females from provisioning alternate nests or preventing them from both ovipositing and provisioning. Our results indicate that both egg-laying and provisioning represent significant costs of reproduction, expressed as differences in productivity but not survival. A trade-off-based model suggests that other components of parental care such as nest initiation may also represent significant costs. Costs of egg production and nest initiation are probably similar for male and female offspring, so that taking them into account leads to a less male-biased expected sex ratio. Mothers compensated only partially for prey removal in terms of the total provisions they gave to individual offspring.

The origin and maintenance of eusociality is a central problem in evolutionary biology1, 2. Eusocial groups contain individuals that forfeit their own reproduction in order to help others reproduce. In facultatively eusocial taxa, offspring can choose whether to found new nests or become helpers in their natal groups. In many facultatively eusocial insects, offspring need continuous care during development, but adult carers have life expectancies shorter than the developmental period3, 4, 5, 6, 7. When a lone foundress dies, her partly reared brood are usually doomed. Here, we show that helpers in a tropical hover wasp (Liostenogaster flavolineata) have an insurance-based advantage over lone foundresses because after a helper dies, most of the brood that she has partly reared will be brought to maturity by surviving nest-mates. After some of the helpers are experimentally removed from a multi-female nest, the reduced group is left with more brood than it would normally rear. We found that larger, more valuable extra brood were reared through to maturity, but not smaller, less valuable brood. Smaller brood may be sacrificed to feed larger brood, and reduced groups probably benefited from increased short-term helper recruitment. Rearing extra brood did not increase adult mortality or brood development time.

A candidate explanation for the evolution of eusociality is that helpers are physiologically constrained such that helping is their only realistic option. We tested this subfertility hypothesis in a species of facultatively eusocial hover wasp (Hymenoptera, Stenogastrinae: Liostenogaster flavolineata) by seeing whether helpers that were forced to nest on their own were able to mature their own eggs. One focal helper was left alone on each of 22 nests, from which all other adult wasps (including the dominant) were permanently removed. After 18 days, all but one of the 19 focal helpers that remained on their nests had ovarian development and insemination status characteristic of dominants, and the majority had probably laid eggs. This was in striking contrast to the reproductive status of other helpers removed from the same nests at the start of the experiment. These results provide convincing experimental evidence that females do not become helpers because of some unconditional physiological constraint. There is currently no unequivocal support for the subfertility hypothesis in facultatively eusocial Hymenoptera lacking morphological castes. Copyright 1999 The Association for the Study of Animal Behaviour.

Recent field experiments suggest that cooperative breeding in vertebrates can be driven by a shortage of breeding territories. We did analogous experiments on facultatively eusocial hover wasps (Stenogastrinae: Liostenogaster flavolineata). We provided nesting opportunities by removing residents from 39 nests within a large aggregation (1995), and by glueing 20 nests obtained from a distant site into a second aggregation (1996). We prevented nest-less floaters from competing for these opportunities in 1995 but not in 1996. In both years, helpers in unmanipulated groups were given opportunities to nest independently without having to incur nest-building costs and with a reduced wait before potential helpers emerged. Helpers visited the nests we provided, but adopted only a small proportion (5% of 111 vacancies created in 1995). Others were adopted by floaters, but a significant proportion of nests were never adopted (nine out of 20 in 1995, seven out of 20 in 1996). Helpers that visited nests did not originate from particular kinds of social group. Nests containing older brood were more likely to be adopted, and adopting females rarely destroyed older brood. A general feature of social insect but not vertebrate life-histories, the long period of offspring dependency relative to the short life expectancy of adult carers, may be a key factor constraining independent nesting.

Nesting tactics of individually marked female solitary wasps, Ammophila sabulosa (L.) (Sphecidae), were studied over two seasons at a Norfolk heathland site. Females provision unicellular nests with 1-5 paralysed caterpillars. The entire nesting cycle requires approximately 8-10 hours of activity and there is no evidence that nesting success depends on female size. Prey are obtained by hunting or are stolen from other females' nests, which are found by searching in the nesting area. Females search on the nesting area and open other females' nests at significantly later times of day and lower temperatures than those at which they are absent from the nesting area (presumably hunting) and provision hunted prey. This may reflect thermal constraints or changes in prey availability or availability of detectable nests. Hunting and searching are mutually exclusive alternatives, but because females frequently switch between them it is difficult to estimate the time costs of either. Brood parasitism is an alternative way in which an A. sabulosa female can provide her larva with food and shelter. After completing her previous nest she must find a new digging site. While searching in the nesting area she may come across another female's nest, which she opens and usually brood parasitizes if it contains prey. It is again difficult to compare the time costs of non-parasitic nesting and parasitism, this time because alternative behaviours are not mutually exclusive - finding digging sites and finding other females' nests both involve searching on the nesting area. Females might find other females' nests opportunistically, or spend time searching for them. My methods and interpretation are compared with those of BROCKMANN et al., who analysed alternative nesting strategies in Sphex ichneumoneus. The alternative behaviours in Sphex are 'digging' and 'entering', two ways of acquiring a burrow. BROCKMANN et al. included nights and periods of poor weather in their time cost measurements, decreasing the likelihood of detecting differences between the net benefits of digging and entering. They also assumed that eggs laid after digging and entering had equal chances of survival, and that a female decides unconditionally to either dig or enter immediately after closing her previous nest. A major problem in both studies is that of pin-pointing when decisions are made and hence defining the strategy set. This is particularly difficult when alternative behaviours are not mutually exclusive.

The nesting behaviour of marked female solitary wasps, Ammophila sabulosa (L.) (Sphecidae), was studied over two seasons at a Norfolk heathland site. Females mass provisioned unicellular nests with 1-5 paralysed caterpillars. Intraspecific parasitism was the major mortality factor before the egg hatched - approximately 28% of eggs were destroyed by parasites. Parasites stole caterpillars to provision their own nests, replaced host eggs with their own eggs (brood parasitism), or simply discarded prey from host nests. Hosts were ineffective at preventing parasitism. Individual females both provisioned their own nests and parasitized those of other females. Parasitism usually occurred soon after the previous activity at the host nest. Parasitism events were clumped at certain nests, which were parasitized up to five times each. Once a nest had been brood parasitized, the prey were significantly more likely to be stolen or discarded than were prey in unparasitized nests, and only 17 % of eggs laid by brood parasites survived. Circumstantial evidence suggests that chemical stimuli are important in detection of host nests by parasites. High levels of intraspecific parasitism have rarely been reported in solitary wasps and bees, and parasitic behaviour is rare and unspecialized in some other Ammophila species. Parasitism by miltogrammine flies was of relatively minor importance in A. sabulosa as in other Ammophila species, probably because provisioning events, during which flies larviposit, are few and well separated in time.

Queues formed by social wasps to inherit the dominant position in the nest are analyzed by using a transient quasi-birth-and-death (QBD) process. We show that the extended nest lifespan due to division of labor between queen and helpers has a big impact on nest productivity.

Temperate-zone socially polymorphic sweat bees (Hymenoptera: Halictidae) are ideal model systems for elucidating the origins of eusociality, a major evolutionary transition. Bees express either social or solitary behaviour in different parts of their range, and social phenotype typically correlates with season length. Despite their obvious utility, however, socially polymorphic sweat bees have received relatively little attention with respect to understanding the origins of eusociality. Lasioglossum (Evylaeus) calceatum is a widespread sweat bee that is thought to be socially polymorphic, with important potential as an experimental model species. We first determined the social phenotype of L. calceatum at three sites located at different latitudes within the UK. We then investigated sociality in detail across two years at the southernmost site. We found that L. calceatum exhibits latitudinal social polymorphism within the UK; bees were solitary at our two northern sites but the majority of nests were social at our southern site. Sociality in the south was characterised by a relatively small mean of two and 3.5 workers per nest in each year, respectively, and a small to medium mean caste-size dimorphism of 6.6 %. Foundresses were smaller in our more northern and high altitude populations. Sociality is clearly less specialised than in some closely related obligately social species but probably more specialied than other polymorphic sweat bees. Our research provides a starting point for future experimental work to investigate mechanisms underlying social polymorphism in L. calceatum.