Hibernation clustering and its consequences for associative nest foundation in<Emphasis Type="Italic"> Polistes dominulus</Emphasis> (Hymenoptera Vespidae)

Various studies indicate that cuticular hydrocarbons (CHCs) act as cues for nestmate recognition in paper wasps and other social insects. A recent study showed that associative nest foundation in Polistes dominulus is mainly performed by foundresses coming from the same locality. In the present study, we induced future foundresses of P. dominulus collected in two different localities to hibernate in the laboratory in aggregates of individuals from the same or different localities. After 2 months of hibernation, foundresses from different localities but from the same experimental cluster did not show any preference to associate, at the time of nest foundation, with individuals coming from the same original locality. The cuticular chemical profiles of individuals from the mixed hibernation clusters were quite similar and significantly different from those of individuals which hibernated with other foundresses from the same locality. These findings suggest that, in this species, mechanisms other than nestmate chemical recognition play a major role in the spring association of gynes during the foundation of a new nest.Communicated by R.F.A. Moritz     

Worker rank, reproductive status and cuticular hydrocarbon signature in the ant, Pachycondyla cf. inversa

Workers in de-queened colonies of the neotropical ant, Pachycondyla cf. inversa, form linear or near-linear dominance hierarchies by violent antennation and biting. In these rank orders, social status and ovarian activity are on average highly correlated. Whereas the presence of a fertile queen appears to be sufficient to prevent workers from laying eggs, fertile workers do not completely control reproduction by their nestmates, suggesting that workers are able to differentiate between an egg-laying queen and an egg-laying worker. Here we show that the composition of cuticular hydrocarbons of egg-laying workers is quantitatively and qualitatively different from that of non-laying workers and resembles the hydrocarbon blend of the queen but does not completely match it. Furthermore, using discriminant analysis, it was possible to distinguish workers with four different classes of ovarian development based only on their cuticular hydrocarbon profiles. Fertility-associated changes in cuticular hydrocarbons may play an important role in the behavioural regulation of reproduction in this ant.     

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.     

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.     

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.     

Social organization and division of labour in colonies of the polistine wasp,Belonogaster petiolata

Summary Foundresses in pre-emergence and post-emergence nests of Belonogaster petiolata were organized into linear dominance hierarchies according to their level of physical aggression towards cofoundresses. The female at the top of the hierarchy became the queen, while foundresses ranked below her became worker-like subordinates. In pre-matrifilial colonies, worker offspring were socially subordinate to both their queen and the subordinate foundresses. Queens of matrifilial colonies retained full social dominance over their workers. Queens were reproductively dominant over subordinates and workers, and laid the majority of, if not all, surviving eggs. Subordinate-laid eggs were invariably discovered and destroyed by the queen through oophagy; workers in pre-matrifilial and matrifilial colonies never laid or ate eggs. Colonies contained a single (and the same) functional queen throughout the pre- and post-emergence periods and were, therefore, long-term monogynous.In both established pre-emergence colonies and post-emergence colonies, virtually all foraging for food and nesting material was performed by the subordinates and workers. More dominant subordinates generally foraged less than low-ranked subordinates. Queens were more active builders than subordinates or workers in pre-matrifilial and matrifilial colonies. Queens enjoyed a surplus in exchanged food loads, while subordinates and workers did not. Among cofoundresses, an advantage in food exchange was also positively associated with dominance rank. Queens solicited larvae for their salivary fluid more often than did subordinates and workers. Among cofoundresses, frequency of adult-larva trophallaxis was positively associated with rank. Queens rested most often in the central zones of the nest containing late instar larvae and pupae, while subordinates and particularly workers spent more time in the peripheral, broodless regions of the nest.This paper is dedicated to the late Professor Leo Pardi, whose pioneering studies of Belonogaster served as a constant source of inspiration and reference for the present work     

Chemical mimicry in the root aphid parasitoid Paralipsis eikoae Yasumatsu (Hymenoptera: Aphidiidae) of the aphid-attending ant Lasius sakagamii Yamauchi & Hayashida (Hymenoptera: Formicidae)

Summary. An aphidiid wasp, Paralipsis eikoae, was associated with both Lasius niger and L. sakagamii attending the wormwood root aphid Sappaphis piri. An L. sakagamii worker was observed carrying a winged female P. eikoae to its nest with its mandible, but it did not kill the wasp. Once accepted by the ants, the wasp often mounted and rubbed against the worker ants and sometimes teased them to regurgitate food to itself. No workers in the colony attacked the wasp. Conspecific foreign workers, however, viciously attacked the wasp when encountered. Gas chromatography-mass spectrometry analyses showed that the accepted wasp had complex cuticular hydrocarbons that were very similar to those of its host ants, whereas the winged wasps collected outside the ant nest showed only a series of n-alkanes. Additionally, the accepted wasp had a hydrocarbon profile closer to that of its host ants than to the conspecific foreign ants. We believe the wasp mimics ant cuticular hydrocarbons to integrate into the ant nest, acquiring the hydrocarbons by mounting and rubbing against the ants. In contrast, the cuticular hydrocarbons of the emerged wasp contained larval and pupal hydrocarbons of L. sakagamii that were also similar to those of L. niger. Both ant species rejected adult workers of the other species but accepted their larvae and pupae. We suggest that the emerged P. eikoae mimics the cuticular hydrocarbons of these Lasius larvae and pupae, which allows P. eikoae to be accepted by both L. sakagamii and L. niger. Received 11 March 1998; accepted 22 July 1998.     

Behavioural evidence for the involvement of Dufour’s gland secretion in nestmate recognition in the social wasp Polistes dominulus (Hymenoptera: Vespidae)

The results of recent studies by our group have suggested that in social wasps of the genus Polistes the Dufour’s gland is involved in kin recognition. In fact, the same hydrocarbons occurring on the cuticle are found in the gland secretion, and in P. dominulus the composition of the glandular secretion is more similar in foundresses belonging to the same colonies than in heterocolonial foundresses. In this study, P. dominulus colonies were experimentally presented with previously treated dead conspecific females. These lures had been deprived of their epicuticular lipids and coated with epicuticular or Dufour’s gland secretion extracts from females hetero- or homocolonial with respect to the tested colonies. The behaviour of the colonies towards these lures indicates that, like the epicuticular lipids, the Dufour’s gland secretion is involved in nestmate recognition. Received: 14 August 1995/Accepted after revision: 5 November 1995     

Local mate competition in the solitary parasitoid wasp <Emphasis Type="Italic">Ooencyrtus kuvanae</Emphasis>

Local mate competition (LMC) occurs when brothers compete with each other for mating opportunities, resulting in selection for a female-biased sex ratio within local groups. If multiple females oviposit in the same patch, their sons compete for mating opportunities with non-brothers. Females, in the presence of other females, should thus produce relatively more sons. Sex ratio theory also predicts a more female-biased sex ratio when ovipositing females are genetically related, and sex-ratio responses to foundress size if it differentially affects fitness gains from sons versus daughters. The mating system of the parasitoid wasp Ooencyrtus kuvanae meets assumptions of LMC. Females insert a single egg into each accessible egg of gypsy moth, Lymantria dispar, host egg masses. Wasps complete development inside host eggs and emerge en masse, as sexually mature adults, resulting in intense competition among brothers. We tested the hypothesis that O. kuvanae exhibits LMC by manipulating the number of wasp foundresses on egg masses with identical numbers of eggs. As predicted by LMC theory, with increasing numbers of wasp foundresses on an egg mass, the proportions of emerging sons increased. In contrast, the presence of a sibling compared to a non-sibling female during oviposition, or the size of a female, did not affect the number or sex ratio of offspring produced. The O. kuvanae system differs from others in that larvae do not compete for local resources and thus do not distort the sex ratio in favor of sons. With no resource competition among O. kuvanae larvae, the sex ratio of emergent son and daughter wasps is due entirely to the sex allocation by ovipositing wasp foundresses on host egg masses.     

Alteration of cuticular hydrocarbon composition affects heterospecific nestmate recognition in the carpenter ant Camponotus fellah

Nestmate recognition is a ubiquitous phenomenon in social insects as a means to prevent entry of undesired individuals aiming at exploiting the rich nest resources. The recognition cues in ants were shown in a few cases to be cuticular hydrocarbons, although there are a quite number of correlated associations. In the present study we modified the cuticular profiles of workers Camponotus fellah hydrocarbons with cuticular washes from a closely related, yet undescribed species, Camponotus sp. Although these sympatric species are morphologically indistinguishable, cuticular washes of C. sp. contain 9,13-dimethylpentacosane and 11,15-dimethylheptacosane that are either absent or occur as traces in C. fellah. In addition, C. sp. contains significantly greater amounts of 3-methylpentacosane than C. fellah workers. The cuticle modification was done solventless in a manner that minimized disruption to the cuticular structure of the ant being modified. Judging from the 3 focal compounds, such treatment added between 20 and 30% of the original amounts present in C. sp. to the treated C. fellah workers. This addition changed consistently the cuticular hydrocarbon profile of the treated ant. Dyadic assays between C. fellah and their nestmates treated with C. sp. cuticular rinses revealed a significantly higher level of aggression compared to non-treated nestmates. There was no aggression between nestmates of C. sp. These results demonstrate that in heterospecific interactions between the two Camponotus species there is a correlation between cuticular hydrocarbons and a nestmate recognition response, albeit not as high as the response of C. fellah to of C. sp. workers. This is consistent with the hypothesis that cuticular hydrocarbons may play a role in nestmate recognition.     

Why is dominance hierarchy age-related in social insects? The relative longevity hypothesis

In temperate regions, older eusocial hymenopteran females with annual life cycles (annual-temperate) tend to dominate younger females, a behavior demonstrated by many Polistes. However, in queenless ponerine ants and primitively eusocial tropical wasps (perennial/tropical), a younger female can be dominant and occasionally takes over from the older, most dominant reproductive female, the alpha. We investigated these patterns using an inclusive fitness model. The most important difference between the above two cases lay in the length of individual life compared with colony life. Colonies dissolve before winter in the annual-temperate case, so the expected future tenure of the replacement alpha is never longer than that of the original alpha. This makes the non-reproductive subordinate tactic more advantageous for individuals that emerge later in the season because of the fitness cost of superseding. The perennial/tropical case does not have a clear upper limit for colony longevity, so the model predicts that late-born younger daughters are more likely to challenge their mother-alpha because of the expected long future tenure of the new alpha compared with the small indirect cost of the mothers reproductive failure. To switch tactics from being a subordinate to being the new alpha is only optimal in some situations. The high mortality rate of subordinate workers does not qualitatively alter the above pattern. The specific sexual production schedule of the colony sometimes affects the optimal behavior of females in the perennial/tropical case, and older individuals can dominate younger ones when the end of current round of sexual production is imminent.Electronic Supplementary Material Supplementary material is available for this article at     

Dominant male red junglefowl (Gallus gallus) test the dominance status of other males

In sexual selection, honest signals are maintained by a variety of mechanisms. In red junglefowl (Gallus gallus), health, condition and social status affect comb size, a well-documented predictor of female choice. The comb size of subordinate male junglefowl appears to be suppressed when in the company of other males. One hypothesis for how social status could affect ornament expression in this way involves punishment of cheaters. Under this scenario, dominant males periodically challenge similar males signalling putative high status. For subordinate males, the risk of fighting a high-ranked male could make it prohibitively costly to develop ornamentation signalling dominance. We asked if dominance signals influenced the direction of aggression by dominant males. To address this issue, we conducted experiments in which 19 dominant-acting, large-combed male junglefowl were allowed to choose to fight one of two opponents. The two potential fight opponents differed in comb size, dominance behaviour, or in both traits. In 15 of 19 trials, dominant-acting males chose to fight large-combed, dominant-acting opponents rather than small-combed, subordinate-acting opponents. This is the first demonstration that aggression of dominant male birds is directed at other males based on the display of an ornament known to be attractive to females. However, males did not discriminate between fight opponents when potential opponents differed in only one of the two status indicators (large-combed males chosen in 11 of 19 trials, dominant-acting males chosen in 10 of 19 trials).     

Behavioral and physiological factors associated with juvenile hormone in Polistes wasp foundresses

Although there is increasing interest in the evolution of endocrine systems, relatively little is known about the factors associated with natural endocrine variation in invertebrates. Here, we assess juvenile hormone (JH) titers among nest-founding queens of the wasp Polistes dominulus over 2 years. We allowed unfamiliar wasps to battle for dominance and examined the relationships between dominance rank, JH, ovarian development, and facial patterns. The relationship between JH-titer and dominance varied across years; there was a stronger relationship between JH-titer and dominance in 2006 than in 2008. Across years, wasps that won dominance contests had facial patterns with more broken black spots than wasps that lost dominance contests. There was no relationship between dominance rank and ovarian development. The individual characteristics associated with JH-titer were also tested; JH-titers were correlated with facial pattern brokenness and ovarian development. This study adds to previous work indicating that P. dominulus facial patterns function as a signal of fighting ability. Furthermore, the correlation between JH-titers and facial patterns parallels previous work on testosterone and vertebrate signals and suggests that links between signals of fighting ability and hormones that mediate fighting ability may be common across taxa. Overall, individual JH-titers vary, though they are typically associated with factors related to individual reproductive success, including dominance, fertility, and facial pattern brokenness. Future studies in additional contexts and taxa will be important to test how and why JH-titers vary.     

Dominance status influences female reproductive strategy in a territorial African rodent <Emphasis Type="Italic">Rhabdomys pumilio</Emphasis>

Dominance status influences the fitness of many mammals. Using African striped mice Rhabdomys pumilio, we tested whether (1) dominant females have greater reproductive success than subordinate females, (2) dominant females influence the reproductive output of subordinate females when they are housed in close proximity, (3) reproductive output of a female changes in response to the dominance status of her neighbours, and (4) whether prolonged association between individuals influences the variance in reproductive success between dominants and subordinates (i.e. the ‘dear enemy’ phenomenon). The size and mass of litters of dominants increased significantly when housed adjacent to subordinates than when housed apart. The litter size and mass of subordinates remained unchanged, although subordinates spent significantly more time with their pups when housed close to dominants than when housed apart; time spent with pups by dominants remained unchanged. Moreover, females modified their reproductive output and behaviour in relation to the dominance status of their neighbours. Following prolonged association, dominants still had greater reproductive success, but now, the time spent with pups decreased in subordinates. We suggest that dominants adopt a strategy to increase the reproductive value of their litter, whereas subordinates adopt a pup defence strategy. These strategies are flexible and are influenced by the dominance status and period of association between neighbours, so that females could maximize their fitness in response to varying social conditions.     

Symmetry, male dominance and female mate preferences in the Iberian rock lizard, Lacerta monticola

Female preference for dominant males is widespread and it is generally assumed that success in male-male competition reflects high quality. However, male dominance is not always attractive to females. Alternatively, relatively symmetric individuals may experience fitness advantages, but it remains to be determined whether males with more symmetrical secondary sexual traits experience advantages in both intra- and intersexual selection. We analysed the factors that determine dominance status in males of the lizard Lacerta monticola, and their relationship to female mate preference, estimated by the attractiveness of males' scents to females. Sexually dimorphic traits of this lizard (head size and femoral pores) appear to be advanced by different selection pressures. Males with relatively higher heads, which give them advantage in intrasexual contests, were more dominant. However, head size was unimportant to females, which preferred to be in areas marked by relatively heavier males, but also by males more symmetric in their counts of left and right femoral pores. Chemicals arising from the femoral pores and other glands might honestly indicate quality (i.e. related to the symmetry levels) of a male to females and may result from intersexual selection. Females may use this information because the only benefit of mate choice to female lizards may be genetic quality. Chemical signals may be more reliable and have a greater importance in sexual selection processes of lizards than has previously been considered.     

The scent of dominance: female field crickets use odour to predict the outcome of male competition

Pheromones may convey information about mate quality and social status. In the field cricket Gryllus integer, females mount the males for copulation, such that males cannot coerce females to mate. We examined whether virgin G. integer females preferred the scent of potentially dominant males to that of subordinate males. First, we collected pheromones by confining males on filter paper. Next, we offered filter paper from each of two size-matched males and control paper to females that had never been exposed to males, and measured the time spent by the female on each kind of paper. Finally, dominance status of the males in each size-matched pair was determined by pitting the two males against one another in agonistic contests. When offered filter paper from subsequently dominant versus subsequently subordinate males, females spent more time on the paper from the dominant male than the subordinate male, and much less time on control paper. Thus, pheromones may inform female G. integer about a male's potential to achieve dominant social status. Male pheromones were also associated with the female's tendency to mount a male. In contrast to cockroaches, where females prefer the scent of subordinate males (presumably to avoid risk of injury), female crickets prefer the scent of potentially dominant males and are more likely than males to wound their mating partners.     

Sexual selection and species recognition in the pygmy swordtail, Xiphophorus pygmaeus: conflicting preferences

Sexual selection and species recognition play important roles in mate choice; however, sexual selection preferences may overlap with traits found in heterospecifics, producing a conflict between sexual selection and species recognition. We examined female preferences in Xiphophorus pygmaeus for male traits that could provide both types of information to determine how females use multiple cues when preferences for these cues would conflict. We also examined X. pygmaeus behavior in the field to determine if females have the opportunity to choose mates. As no male-male competition was observed in the field, and females occasionally chased males from feeding areas, females apparently have the opportunity to exercise mate choice in their natural habitat. In the laboratory, female X. pygmaeus used body size as a sexual selection cue, preferring large heterospecifics (X. cortezi) to small conspecifics. Females also preferred barless X. cortezi over barred X. cortezi when males were size matched. Because X. pygmaeus males do not have bars, this preference suggests that X. pygmaeus females use vertical bars in species recognition, and that large body size and vertical bars are conflicting cues. However, X. pygmaeus females did not have a preference for males of either species when sexual selection and species recognition cues were presented concurrently. This result was surprising, because preferences for species recognition cues are often assumed to be stronger than sexual selection cues. We suggest that females may be using additional species-specific cues in mate choice to prevent hybridization.     

Dominance orders in the ponerine ant Pachycondyla apicalis (Hymenoptera,Formicidae)

Summary Intracolony aggression among workers of the neotropical ponerine ant species Pachycondyla apicalis leads to dominance orders. Antagonistic interactions can entail either overt physical attacks with the subordinate individual often exhibiting a submissive posture or the robbing and destruction of eggs laid by nestmates. The single queen, however, was never observed either attacking or being attacked by any colony member. The hierarchical structure among workers consists of one dominant individual and several subordinates; the relationships among subordinate workers are unclear, however. We report for the first time a natural (nonmanipulated) change in the social status of individuals within an ant dominance order. Dominant workers usually had better developed ovaries, laid more eggs and were more frequently observed attending the egg pile than subordinate individuals. This pattern became even more striking when the queen was excluded from the colony. These results indicate that workers of P. apicalis lay eggs even in the presence of the queen. It is possible that some of these haploid eggs may develop into males.Offprint requests to: B. Holldobler     

Gonad morphology, sexual development, and colony composition in the obligate coral-dwelling damselfish Dascyllus aruanus

Gonad morphology and colony composition support the existing supposition that the obligate coral-dwelling damselfish Dascyllus aruanus has a protogynous hermaphroditic sexual pattern. Adults had either an active ovary containing vitellogenic oocytes, an ovotestis, or a spermiated testis and were classified as adult female, hermaphrodite, or adult male, respectively. Among individuals having male function, the testis (or testis portion of the ovotestis) takes the form of an unrestricted spermatogonial lobular testis. Among hermaphrodites having an ovotestis, a small proportion of individuals had a gonad in which both the ovarian and testicular portions were inactive (inactive hermaphrodites), whereas the majority had a predominantly testicular ovotestis that contained spermatozoa (male-active hermaphrodites). The size range of individuals within gonadal classes indicates that all D. aruanus first develop an ovariform gonad. Some individuals then undergo ovarian maturation to become adult females while others develop testicular tissue to form an ovotestis and become male-active hermaphrodites. Subsequently, progressive loss of ovarian tissue results in the development of a secondary testis from an ovotestis with the retention of a residual, afunctional lumen among adult males. The wide size range of individuals having an ovotestis suggests that some hermaphrodites function as adult females before developing testicular tissue while other individuals do not pass through an adult female stage. If this is the case, D. aruanus exhibits a diandric protogynous hermaphroditic sexual pattern. The apparent prolonged retention of an ovotestis with both healthy oocytes and an ovarian-type lumen in a spermiated ovotestis, as well as a functional sex ratio of 1:1 for adult females:adult males plus male-active hermaphrodites also raises the possibility that D. aruanus may be capable of bidirectional sex change during the hermaphroditic stage. Such a capability would be highly adaptive for a species having limited mobility and unpredictable recruitment of new colony members resulting in unpredictable mating opportunities.     

Effects of male dominance and courtship display on female choice in the ring-necked pheasant

Male traits and behaviours acting in mate choice and intrasexual competition are expected to be congruent. When studying their evolution, this often makes it difficult to differentiate between these two components of sexual selection. Studies are therefore needed on mate choice in conjunction with the role of displays and dominance. We present the results from two experiments conducted to investigate the effects of male dominance and courtship displays on female choice in the ring-necked pheasant, Phasianus colchicus, controlling for differences in morphological male traits. We found: (1) different courtship behaviours had different effects on female choice: females were mainly attracted by the feeding courtship behaviour, while another courtship display (the lateral display) was effective in producing the copulation-acceptance response by the females; (2) subordinate males performed the courtship behaviour before females less frequently than dominant males, and females reinforced intrasexual selection by choosing dominant males, and (3) subordinate males in visual contact with a dominant became less attractive to females. The results support the idea (armament-ornament model) that female pheasants may benefit from using traits selected in male-male competition as clues for mate choice. Received: 23 October 1997 / Accepted after revision: 7 October 1998