Aggression and resource sharing among foundresses in the social wasp Polistes dominulus: testing transactional theories of conflict

Tug-of-war models of within-group conflict predict that the frequency of conflict will be positively related to the degree of reproductive sharing within the group; in contrast, a negative relationship supports transactional models, in which reproductive payments among group members limit the degree of within-group selfishness. We tested predictions of the tug-of-war and transactional models by examining cofoundress interactions during the founding (preworker) phase of colony development in 30 naturally nesting colonies of the paper wasp Polistes dominulus. We found that the mean rate of foundress aggression and the mean probability of food sharing were significantly negatively associated, which supports the prediction of the transactional, not the tug-of-war model. Further, cofoundress aggression significantly increased over the founding phase (independently of temperature), while the fraction of aggression initiated by the dominant (alpha) foundress significantly decreased over this period. We show that both of these results are predicted by the transactional model of within-group conflict. Interestingly, the alpha’s rate of aggression was significantly positively temperature dependent, while the beta’s was not. This indicates that the alpha’s aggression level may often be near her physiological maximum, while the beta’s aggression is limi- ted by other factors, contradicting the prediction of the tug-of-war model. Moreover, the alpha’s aggression was significantly positively temperature dependent only in the second half of the founding period, as predicted by the transactional model since this is when there is least reproductive sharing. Finally, our results indicate that the alpha’s level of aggression depends on the resources controlled by the beta. Received: 18 January 2000 / Revised: 19 June 2000 / Accepted: 24 June 2000     

Multiple reproductive strategies in a tropical hover wasp

Reproductive skew theory has been an important component of efforts to design a unifying theory of social evolution, as it aims to explain patterns of reproductive partitioning in animal societies as a function of relatedness, group productivity, fighting ability and ecological constraints on independent reproduction. However, empirical tests of the theory have often provided ambiguous or non-conclusive results, assumptions behind alternative models have rarely been tested, and theoretical elaborations have shown the limitations of the reproductive skew approach. Here we analyse a relatively large sample of colonies of the Stenogastrine wasp Parischnogaster mellyi with a powerful set of DNA microsatellite markers. We show that various apparently stable forms of social organisation co-exist in a single population, and that sharing of reproduction between related and unrelated egg-laying females occurs in some of the nests. Present reproductive skew theory appears to be at best partly sufficient to account for the observed complexity of social organisation. The observed patterns of colony composition and reproductive sharing are weakly consistent with the hypothesis of reproductive transactions, while they can more parsimoniously be explained by the life-history characteristics of the species.Communicated by R.F.A. Moritz     

Cuticular hydrocarbons and reproductive status in the social wasp Polistes dominulus

Most species of social insect are characterized by a reproductive division of labor among morphologically specialized individuals. In contrast, there exist many species where all individuals are morphologically identical and dominance relationships determine which individuals mate and/or reproduce. In newly founded multiple-foundress associations of the social wasp Polistes dominulus, foundresses establish dominance hierarchies where the top-ranked (alpha) female monopolizes egg laying. The possibility that chemical cues are used for recognition of egg-laying individuals has not been explored in this wasp. Using non-destructive techniques, we examined the relationship between ovarian activity and the proportions of cuticular hydrocarbons of three female types (dominant and subordinate foundresses and workers) in 11 colonies. Immediately after nest foundation, no differences were found between alpha and subordinate females. In contrast, at worker emergence, alpha females were statistically distinguishable from both subordinates and workers. We experimentally removed the alpha female in 5 of the original nests and reanalyzed hydrocarbon proportions of the new dominant individual. Replacement individuals were all found to acquire a cuticular signature characteristic of the alpha female. This suggests that cuticular hydrocarbons are used as cues of ovarian activity in P. dominulus, and we discuss our results in terms of a switch from behavioral dominance to chemical signaling in this wasp.     

Timing matters when assessing dominance and chemical signatures in the paper wasp Polistes dominulus

A recent study (Izzo et al., Behav Ecol Sociobiol 64: 857–864, 2010) reported that cuticular hydrocarbons (CHCs) correlate with fertility, not dominance, in the paper wasp Polistes dominulus thus contradicting the results of recent investigations which concluded that social dominance is the main determinant for CHC signatures in this species. We suggest here that different forms of dominance in the pre-nesting and post-nesting phases caused the apparently contradictory results. Thus the assumption that dominance behaviour in the pre-nesting stage is synonymous with dominance after colony foundation is incorrect. We provide standardised definitions for forms of “dominance” observed in the P. dominulus life cycle to avoid apparent discrepancies in the future among studies dealing with the same topics in different annual stages.     

Egg mimicry and allopaternal care: two mate-attracting tactics by which nesting striped darter (Etheostoma virgatum) males enhance reproductive success

In a variety of fish species with paternal care of offspring, females prefer to spawn in nests that already contain eggs. This female preference has been hypothesized to explain egg thievery in male sticklebacks, allopaternal care of eggs in minnows, and the evolution of egg-mimicking body features in male cichlids and darters. Here we employ microsatellite-based parentage analyses to evaluate the reproductive success of striped darter (Etheostoma virgatum) males that appear to utilize two of these functionally related tactics to entice females to spawn in their nests. In an isolated population (Clear Creek, Ky.), we observed that breeding males develop conspicuous white spots on their pectoral fins. If these spots are egg mimics, as we suspect, then this represents the fourth independent evolutionary origin of egg mimicry documented to date in darters, the first based on pigmentation (as opposed to physical structures), and the first in which the egg mimics vary greatly in number among males. From direct counts of microsatellite genotypes in clutches of embryos, at least 3.8 females contributed to the progeny within a typical nest, and females tended to spawn preferentially with males that were larger and displayed more egg-mimic spots. In another population (Hurricane Creek, Tenn.) without egg mimics, the multi-locus genetic data document that allopaternal care is common, especially among the smallest males who sometimes tend nests containing their own as well as an earlier sire's offspring. Thus, these foster males had adopted egg-containing nests and then successfully spawned with subsequent females. Overall, the genetic data on paternity and maternity, in conjunction with field observations, suggest that egg mimicry and allopaternal care are two mate-attracting reproductive tactics employed by striped darter males to exploit female preferences for spawning in nests with 'eggs'. Electronic Publication     

Reproductive success in a low skew, communal breeding mammal: the banded mongoose, Mungos mungo

In most cooperatively breeding species, reproduction is monopolised by a subset of group members. However, in some species most or all individuals breed. The factors that affect reproductive success in such species are vital to understanding why multiple females breed. A key issue is whether or not the presence of other breeders is costly to an individual’s reproductive success. This study examines the factors that affect the post-parturition component of reproductive success in groups of communal-breeding banded mongoose (Mungos mungo), where up to ten females breed together. Per-litter reproductive success was low (only 18% of pups survived from birth to independence). Whilst singular breeding was wholly unsuccessful, there were costs associated with breeding in the presence of increasing numbers of other females and in large groups. Synchronisation of parturition increased litter success, probably because it minimises the opportunity for infanticide or decreases competitive asymmetry between pups born to different females. There was no evidence of inbreeding depression, and reproductive success was generally higher in litters where females only had access to related males within their group. I conclude that communal breeding in female banded mongooses represents a compromise between the benefits of group-living and communal pup care on the one hand, and competition between females to maximise their personal reproductive success on the other. Such conflicts are likely to occur in most communal breeding species. Whilst communal breeding systems are generally considered egalitarian, negative effects of co-breeders on individual reproductive success is still an issue.     

Biased parental investment and reproductive success in Gabbra pastoralists

Demographic data from 848 Gabbra households are used to examine the relationships between herd size and reproductive success in relation to sex, in a traditional, pastoralist population. The number of camels in the household herd has a significant positive effect on the reproductive success of both men and women, although the effect of wealth is greater for men, as predicted from evolutionary theory. The greater the number of elder brothers a man has, the lower his reproductive success, as a result of a smaller initial herd and a later age at marriage. This is not true for women –number of elder sisters does not have a measurable effect on a woman’s fertility, although it does have a small, negative effect on the size of her dowry. These results are interpreted as competition between same-sex siblings for parental investment, in the form of their father’s herd, which is more intense between sons than daughters as parental investments are greatest in males. Received: 30 June 1995/Accepted after revision: 23 October 1995     

Male reproductive success in free-ranging feral horses

In the social organization of feral horses, adult males compete to monopolize groups or bands of females, sometimes called harems. Alternative male strategies are to remain alone or with other bachelors or, less commonly, to accept subordinate status within a harem. The hypothesis that dominant harem stallion status confers a reproductive advantage was tested in free-ranging feral horses. The presence of foals in harems headed by vasectomized (VSX) versus intact stallions was used to assess the ability of these stallions to control reproduction in their harems. Of harems headed by VSX stallions, 17 and 33% contained foals during years 2 and 3 post-treatment, respectively. In contrast, 86 and 80% of harems headed by non-VSX stallions contained foals in those years. Acquisition of pregnant mares appeared more likely than sneak copulations by bachelor stallions to account for foals in harems with a single stallion. However, most foals were born into harems that included a subordinate stallion, an occurrence that was undoubtedly exacerbated by the extended breeding season resulting from the sterility of the harem stallion. Thus, in comparing alternative reproductive tactics, bachelors appeared less successful than subordinate stallions within a harem. However, the highest reproductive success was achieved by the harem stallion, further demonstrating that alternative tactics are not equally profitable. Received: 13 July 1999 / Received in revised form: 24 July 1999 / Accepted: 24 July 1999     

Philopatry and mother-daughter associations in bushy-tailed woodrats: space use and reproductive success

Social groups in mammals are often based on overlapping generations of philopatric females, yet few researchers have examined fitness consequences of associations between females and their daughters. We examined survival and reproductive consequences of sharing a natal site with one's mother or daughter in wild bushy-tailed woodrats (Neotoma cinerea). Bushy-tailed woodrats reside on discrete rocky outcrops, and females tend to spend their entire reproductive life on their natal outcrop. Mothers and daughters that co-existed on the natal outcrop were closely associated in space. Juvenile females were more likely to survive their first winter to become breeding yearlings at their natal site if their mother was also present at the natal site during the winter. Juveniles survived equally well with or without their mother where densities of adult females were experimentally reduced, suggesting that females enhance their daughter's survival by facilitating their access to limited resources. Yearling females exhibited no reproductive costs from association with their mothers at the natal site during the breeding season; in fact, yearlings that bred at the natal site simultaneously with their adult mothers tended (P = 0.07) to experience fewer reproductive failures than did those that bred in the absence of their mothers. Mothers did not experience any detectable costs when sharing a natal site with a daughter. We conclude that bushy-tailed woodrats exhibit stable, fitness-enhancing associations among mothers and their philopatric daughters.     

Cuticular hydrocarbons correlate with fertility, not dominance, in a paper wasp, Polistes dominulus

Cuticular hydrocarbons (CHCs) are information-rich signals in social insects that coordinate behaviors within nests. However, in some taxa, the precise information conveyed by CHCs is poorly understood. In particular, there is a debate over whether CHCs convey information about their bearer's dominance or fertility. Distinguishing between dominance and fertility signaling is difficult because fertility and rank are frequently correlated within social insect colonies. This study disentangles those relationships by examining CHCs of Polistes dominulus paper wasps during the early nest-founding stage before dominance and fertility become correlated. First, we confirm that dominance and fertility are not associated in early spring foundresses. Then we show that CHCs are more strongly associated with fertility than dominance. There was no relationship between cuticular hydrocarbon profiles and a wasp’s ability to dominate rivals through aggression, suggesting that CHCs are unlikely to provide information about dominance. However, there was a significant correlation between ovarian development and the CHC profile, suggesting that CHCs could convey valuable information about their bearer's fertility. Furthermore, our data provide a potential mechanism for chemical signaling of fecundity, as there is a relationship between endogenous juvenile hormone titer (a gonadotropin), degree of ovarian development, and the CHC profile. Hormonal regulation of CHC profile expression offers a physiological mechanism to coordinate behavior, physical state, and signal expression.     

Intraspecific comb usurpation in the social wasp Polistes fuscatus

Summary Incidents of usurpation were observed in colonies of Polistes fuscatus nesting on farm buildings (1977–79) and in nestboxes (1980–84) in Johnson County, Iowa, USA. Most usurpations (84.8%) occurred in the latter half of the preworker phase of the colony cycle, which coincided with periods of high predation of combs by vertebrates. Usurpers were probably displaced single foundresses which did not join neighbors or refound colonies after comb loss. Most (89–100%) usurpers of known relatedness to the foundresses they replaced were cousins or less related to them. Usurpation was a significant source of nest loss (19.6%) among single foundresses, but was rare (2.2%) in multiple-foundress colonies and colonies with workers (3.5%). Usurpers often destroyed younger brood (eggs and larvae in instars 1–3) in host colonies, while older larvae and pupae were usually spared. Brood destruction was more pronounced in more advanced host combs. Usurper survivorship after workers eclosed was lower than than of queenright single foundresses (61.5% vs 87.0%). Reproductive success by usurpers was less than that of queenright single foundresses, but greater than that of foundresses which initiated colonies late in the preworker colony cycle.     

Comparative tests of reproductive skew in male primates: the roles of demographic factors and incomplete control

Reproductive skew models have been proposed as a unifying framework for understanding animal social systems, but few studies have investigated reproductive skew in a broad evolutionary context. We compiled data on the distribution of mating among males for 31 species of primates and calculated skew indices for each study. We analyzed the determinants of mating skew with phylogenetic comparative methods to investigate two models from reproductive skew theory, the concession model and the tug-of-war model. Mating skew decreased as the number of males increased in multimale groups, suggesting that monopolization of females becomes more difficult when there are more rivals, and therefore supporting the tug-of-war model. We predicted that single males are unable to monopolize receptive females as overlap in female receptivity increases (estrous synchrony) and, as a result, that mating skew decreases. However, we did not find any evidence for a link between female estrous synchrony and male mating skew. Finally, the concession model predicts high skew in male philopatric species relative to species in which males disperse, yet our measures of mating skew showed no significant associations with qualitative scores of male dispersal. More definitive tests of the concession model will require more quantitative measures of relatedness, which are presently unavailable for most primate species in our study. Overall, our results provide support for the tug-of-war model in primates, and the approach developed here can be applied to study comparative patterns of skew in other biological systems.Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible to authorized users.     

Control of reproduction in social insect colonies: individual and collective relatedness preferences in the paper wasp, Polistes annularis

Social insect colonies often have one or a few queens. How these queens maintain their reproductive monopoly, when other colony members could gain by sharing in the reproduction, is not generally known. DNA microsatellite genotyping is used to determine reproductive interests of various classes of colony members in the paper wasp, Polistes annularis. The relatedness estimates show that the best outcome for most individuals is to be the reproductive egg-layer. For workers, this depends on the sex of offspring: they should prefer to lay their own male eggs, but are indifferent if the queen lays the female eggs. The next-best choice is usually to support the current queen. As a rule, subordinates and workers should prefer the current queen to reproduce over other candidates (though subordinates have no strong preference for the queen over other subordinates, and workers may prefer other workers as a source of male eggs). This result supports the theory that reproductive monopoly stems from the collective preferences of non-reproductives, who suppress each other in favor of the queen. However, we reject the general hypothesis of collective worker control in this species because its predictions about who should succeed after the death of the present queen are not upheld. The first successor is a subordinate foundress even though workers should generally prefer a worker successor. If all foundresses have died, an older worker succeeds as queen, in spite of a collective worker preference for a young worker. The results support the previous suggestion that age serves as a conventional cue serving to reduce conflict over queen succession. Received: 3 May 1996 / Accepted after revision: 22 September 1996     

A test of reproductive skew models in a field population of a multiple-queen ant

Determining the evolutionary basis of variation in reproductive skew (degree of sharing of reproduction among coexisting individuals) is an important task both because skew varies widely across social taxa and because testing models of skew evolution permits tests of kin selection theory. Using parentage analyses based on microsatellite markers, we measured skew among female eggs (n=32.3 eggs per colony, range=20–68) in 17 polygynous colonies from a UK field population of the ant Leptothorax acervorum. We used skew among eggs as our principal measure of skew because of the high degree of queen turnover in the study population. Queens within colonies did not make significantly unequal contributions to queen and worker adult or pupal offspring, indicating that skew among female eggs reflected skew among daughter queens. On average, both skew among female eggs (measured by the B index) and queen–queen relatedness proved to be low (means±SE=0.06±0.02 and 0.28±0.08, respectively). However, contrary to current skew models, there was no significant association of skew with either relatedness or worker number (used as a measure of productivity). In L. acervorum, predictions of the concession model of skew may hold between but not within populations because queens are unable to assess their relatedness to other queens within colonies. Additional phenomena that may help maintain low skew in the study population include indiscriminate infanticide in the form of egg cannibalism and split sex ratios that penalize reproductive monopoly by single queens within polygynous colonies.     

Biparental investment and reproductive success in a subsocial desert beetle: the role of maternal effort

Parastizopus armaticeps is a nocturnal subsocial detritivorous desert tenebrionid that produces very few offspring per brood. The two environmental factors that constrain reproduction, rapid sand desiccation rate and food scarcity, are countered by biparental effort. Males dig and extend breeding burrows, maintaining their moisture level; females forage on the surface at night for high-quality detritus, the larval food. This was shown to be a scarce and unpredictable resource for which there is high competition. When food was supplemented in a field experiment, offspring number and survivorship doubled and burrow failure due to desiccation dropped from approximately half, the typical failure rate for unsupplemented burrows, to zero. Food supplementation did not, however, increase larval foodstore size and there was no difference in the size of the offspring produced. Supplemented females reallocated their time, foraging less and digging more with the male. This change in maternal behaviour patterns resulted in deeper burrows which remained moist longer, thus extending the larval production period. Female foraging efficiency, particularly food retrieval speed, determined how much time females could allocate to digging, consequently increasing the reproductive success of the pair. Burrow depth and sand moisture level at the burrow base were the major correlates of reproductive success, but the scarcity and unpredictability of high-quality food on the surface and the competition for this resource influenced the number of offspring indirectly through their effect on female behaviour. Received: 29 November 1996 / Accepted after revision: 7 December 1997     

Influence of insect hormones on the establishment of dominance hierarchies among foundresses of the paper wasp,Polistes gallicus

Summary Laboratory studies on overwintered foundresses of Polistes gallicus show that differences in the endocrine activity are mainly responsible for achieving the dominance rank. Females that became dominant had either larger corpora allata or more developed ovaries than subordinate females. Body size did not contribute to dominance rank. Since a correlation exists between the length of terminal oocytes and the ecdysteroid titre in haemolymph as well as between the volume of corpora allata and the synthesis of juvenile hormone, dominant behaviour is thought to depend upon an elevated hormone titre in haemolymph. Injections of juvenile hormone (JHI) and 20-hydroxyecdysone, separately and simultaneously, significantly increased the probability that the treated female would be the dominant female of a test pair. After a hierarchy has been established, endocrine activity in subordinate foundresses is inhibited by the dominant foundress that then monopolizes reproduction.     

Mating patterns and reproductive success in the bushy-tailed woodrat (Neotoma cinerea), as revealed by DNA fingerprinting

The mating patterns and reproductive success of the bushy-tailed woodrat (Neotoma cinerea) were investigated over a 3-year period (1992–1994) using DNA fingerprinting. Paternity was determined by genetic analysis of 58 juveniles of known maternity from 35 litters. Analysis of DNA fingerprints revealed that all offspring within a litter were fathered by a single male; the statistical probability of detecting multiple males mating with a female was high, indicating that multiple paternity would have been detected had it occurred. However, individual males did not father more than one litter from a given female either within or between years. At least 75% of females and 57% of males successfully produced offspring each year. The finding that all littermates are first-order relatives may contribute to the high level of female cooperation in this species. Received: 28 May 1997 / Accepted after revision: 22 March 1998