Conserved male-specific cuticular hydrocarbon patterns in the trap-jaw ant Odontomachus brunneus

Cuticular hydrocarbons have been identified as the source of sex-recognition signals for many insects, but for social insects, specifically ants, cuticular hydrocarbon profiles of males are often ignored. This study reports male-specific cuticular hydrocarbon patterns for the trap-jaw ant Odontomachus brunneus. Analysis of samples from four Florida populations demonstrated that male-specific overabundance of four hydrocarbons is conserved across populations despite population-level divergence of the remainder of the profile. In addition, hydrocarbon patterns unique to adult males were not present on the cuticle of final instar male larvae, indicating that male-specific profiles arise late in development. The pattern of an abundant subset of conserved cuticular hydrocarbons characteristic of males across divergent populations was compared to earlier findings of the conservation of fertility signals of females across these same populations.     

Queen life-span and total reproductive success are positively associated in the ant Cardiocondyla cf. kagutsuchi

While reproduction and longevity are negatively correlated in a large number of animals, this fundamental trade-off appears to be absent from social insect queens. Here, we examine the distribution of the life-spans of queens and their mates and the age-trajectory of reproduction in experimental colonies of the Pacific tramp ant Cardiocondyla cf. kagutsuchi. Queens of Cardiocondyla are generally short-lived (0.5–2 years), which allows determining their lifetime reproductive success and establishing mortality tables. We show that the queen’s total number of sexual offspring is positively associated with its life-span. Regular counts of eggs, larvae, and pupae suggest that more fecund queens live longer than less fecund queens and that an early onset of sexual production does not negatively affect the queen’s life-span. This corroborates the view that the costs of reproduction are not born by the queen itself. The number of eggs present in colonies increased with queen’s age until shortly before death, indicating negligible reproductive senescence. Sex ratios were strongly affected by the origin of queens but neither by colony size nor total productivity. Several queens produced only males late in their lives, suggesting the occurrence of sperm depletion. Male life-span was not correlated with any other studied trait. Our study shows that the ant genus Cardiocondyla provides a suitable model system to investigate the evolution of life-span in social insect queens and males.     

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.     

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     

Co-evolution-driven cuticular hydrocarbon variation between the slave-making ant Rossomyrmex minuchae and its host Proformica longiseta (Hymenoptera: Formicidae)

Summary. Proformica longiseta exists as two populations in the Sierra Nevada Mountains in Spain, only one of which is parasitized by the slave-maker ant Rossomyrmex minuchae. To investigate the possible effect of co-evolutionary pressures on cuticular hydrocarbon (CHC) profiles (the presumed nestmate recognition cues), we performed a comparative analysis of the CHC of R. minuchae and P. longiseta colonies from both the allopatric and sympatric populations; the latter includes samples of enslaved as well as free-living workers. Discriminant analyses based on these chemical profiles showed two clear profile groups: the first comprised R. minuchae and both enslaved and free-living P. longiseta from the sympatric population; and the second the allopatric P. longiseta workers. As expected, the profiles of the two sympatric P. longiseta groups (enslaved and free-living) were distinct; but, interestingly, those of the enslaved P. longiseta and its parasite R. minuchae were also distinguishable. This indicates that despite their cohabitation each species maintains its own chemical identity. Profile similarity between the sympatric free-living P. longiseta and its parasite may explain the lower than expected aggression observed during raids. We further speculate that in view of the differences between the sympatric and allopatric population of P. longiseta, co-evolutionary pressures have driven changes in the profile of the former to better match that of its parasite R. minuchae. Such an adjustment may have enabled nests of the sympatric P. longiseta to endure multiple raids by the parasite (due to the reduced aggression) and thus to continue to reproduce despite the damage inflicted by the raids.     

The front basitarsal brush in Pachycondyla apicalis and its role in hydrocarbon circulation

Summary. Worker postpharyngeal gland (PPG) chemistry of Pachycondyla apicalis constitutes a blend of hydrocarbons including alkanes, alkenes, and alkadienes, with 5,9-pentacosadiene, 5,9-heptacosadiene and 5,9-nonacosadiene as the major products as well as fatty acids. There is a large congruency between the PPG and cuticular hydrocarbons, however the latter had higher proportion of tricosane and pentacosane, but only traces of fatty acids. These ants have a well-developed cuticular hair brush on the front legs that they frequently pass through the mouth while grooming. Since these brushes have higher amounts of hydrocarbons than either the middle or hind legs, we hypothesized that they are involved in hydrocarbon circulation. We assessed the role of the front basitarsal brush in hydrocarbon circulation by isolating them from the buccal cavity, either by restraining the front legs or by blocking the mouth with glue. In both experiments, preventing contacts between the basitarsal brushes and the mouth for 24 hrs resulted in a considerable increase in hydrocarbon amounts in the brushes with a parallel decrease in the PPG. When the ants were kept alive for an additional 24 hrs after their legs were freed, hydrocarbon amounts in both the PPG and the front legs returned to normal. Analyses of crop content revealed the same hydrocarbons profile as in the PPG, but their amounts were very variable and there were no differences in amounts between the treatment and the control. We assume that part of the PPG hydrocarbons are cleared via the alimentary canal. Grooming behavior of ants restrained for 24 hrs and then freed was followed for four hours thereafter. There was a significant increase in total grooming during the first third and fourth hour, but not during the second. Grooming involving the front legs-mouth, and front legs-antennae, and the front and hind legs was especially intensive during the first hour. Grooming involving the hind legs - body was more intensive in the treated ants during the third hour following their release. We conclude that the front basitarsal brush is a site of hydrocarbon accumulation and that the PPG receives frequent charging during self-grooming.     

Worker population structure and gyne production in the ant Myrmica

Summary Wild colonies of Myrmica rubra L. were sampled and cultured in the laboratory to assess their reproductive capability. Correlation analysis was then used to relate the reproductive performance to a number of worker population characteristics. This revealed an association between the magnitude of gyne production and the proportion of nurses in the population. The absence of big workers also increased gyne production.Artificially composed cultures were then made and used to confirm greater resistance of nurses and small workers to queen domination.It is suggested that rapidly growing colonies will have higher proportions of young nurse workers refractory to queens and able to deliver the food necessary for gyne production. Such colonies could optimise their reproduction in this way.     

Dominance rank,female reproductive synchrony,and male reproductive skew in wild Assamese macaques

In groups with multiple males, direct mate competition may select for the evolution of dominance hierarchies that sort males into a queue for access to fertile females. The priority-of-access (PoA) model proposed by Altmann in 1962 makes explicit predictions about the resulting paternity distribution based on an interaction between male dominance rank and the overlap of female receptive phases. Here, we investigated whether the logic of the PoA model predicted the distribution of paternity across ranks in a seasonal breeder with high reproductive synchrony over six consecutive mating seasons. We studied 18 males that resided in a group of wild Assamese macaques (Macaca assamensis) in their natural habitat at Phu Khieo Wildlife Sanctuary, Thailand, between 2006 and 2011 with 5 to 13 conceptions per season. We assessed whether mate guarding increased paternity success, described “short-term” deviations from predicted paternity distribution, and examined how these are related to the number of competitors and fertile females. We determined genetic paternity of 43 (93 %) offspring born into the study group and found reproductive skew to be relatively low with 29 % alpha male paternity in accordance with the high degree of female reproductive synchrony observed. Short-term deviations from expected paternity distribution over ranks were not explained by the number of resident males or the number of conceiving females or their interaction. Within the limits of this study, these results suggest that even if males cannot discern female fertile phases, if reproduction is seasonal, and if reproductive synchrony is high, males may also compete directly over access to females.     

Dominance rank,resource availability,and reproductive maturation in female savanna baboons

Summary Ten years of data collected from a population of savanna baboons, Papio cynocephalus anubis, residing near Gilgil, Kenya were analyzed to ascertain the extent to which social and ecological factors influence reproductive maturation in females. First sexual swelling occurred at an average age of 4.79 years and first birth occurred at an average age of 6.92 years. Age at first menses was significantly correlated with age at first sexual swelling, but age at first sexual swelling was not a good predictor of age at first birth. The amount of rainfall in the 6 months preceding first sexual swelling and resource availability were significantly correlated with age at first sexual swelling. When ecological factors were taken into account, dominant females had an earlier age at onset of puberty, but not an earlier age at first birth, than did subordinate females. We suggest that nutritional and social stress operate at the same physiological level to disrupt GnRH pulsatility and retard reproductive maturation in some females. Given that socioecological variables modify the timing of life history events related to fitness in female baboons, the task for the future is to unravel how socioecological factors influence different life history components and generate variation in lifetime reproductive success. Correspondence to: F.B. Bercovitch     

Worker reproductive parasitism and drift in the western honeybee Apis mellifera

When a honeybee (Apis spp.) colony loses its queen and is unable to rear a new one, some of the workers activate their ovaries and produce eggs. When a colony has a queen (i.e., it is queenright) almost all worker-laid eggs are eaten, but when hopelessly queenless, the workers become more tolerant of worker-laid eggs and rear some of them to adult drones. This increased tolerance renders a queenless colony vulnerable to worker reproductive parasitism, wherein unrelated workers enter the colony and lay eggs. Here, we show that the proportion of unrelated (non-natal) workers significantly decreases after an Apis mellifera colony becomes queenless. The remaining non-natal workers are as likely to have activated ovaries as natal workers, yet they produce more eggs than natal workers, resulting in significantly higher reproductive success for non-natal workers. In a second experiment, we provided queenless and queenright workers with a choice to remain in their own colony or to join a queenless or queenright colony nearby. The experiment was set up such that worker movement was unlikely to be due to simple orientation errors. Very few workers joined another colony, and there was no preference for workers to drift into or out of queenless or queenright colonies, in accordance with the proportion of non-natal workers declining significantly after becoming queenless in the first experiment.     

Behavioral ecology of the neotropical termite-hunting ant Pachycondyla (= Termitopone) marginata: colony founding,group-raiding and migratory patterns

This study provides the first detailed field account of colony founding, group-raiding and migratory habits in the neotropical termite-hunting ant rPachycondyla marginata, in a semi-deciduous forest in south-east Brazil. New colonies can originate by haplometrosis, pleometrosis, or colony fission. Incipient colonies with multiple foundresses persisted longer in the field, and most excavated nests contained more than one dealated female. A total of 202 group raids by P. marginata were registered, and in all cases the raided termite species was Neocapritermes opacus. Nearly 20% of the workers within a colony engage with raiding activity. Colonies of P. marginata hunt for termites approximately every 2–3 weeks, and group-raids may last for more than 24 h. Target termite nests are up to 38 m from the ant colony, and occasionally two nests are simultaneously raided by one ant colony. Raiding ants carry 1 or 2 paralysed prey, and nearly 1600 termites can be captured during a 9-h raid. Migration by P. marginata colonies lasted over 2 days and covered distances of 2-97 m (n = 48). Average residence time at a given location was 150 days. Three basic migratory patterns were noted: colony fission (only part of the colony moves), long-distance migrations, and short-distance migrations. Both raiding and migratory activities appeared to be strongly affected by seasonal factors. The group raiding and migratory patterns of P. marginata are compared with other ant taxa with similar habits. It is concluded that P. marginata presents a rudimentary form of the so-called army ant behavior, which is highly developed in the subfamilies Dorylinae and Ecitoninae. The extremely specialized diet of P. marginata and the associated high costs of migration are features likely to prevent it from evolving a full army ant life pattern.     

Experimental evidence that workers recognize reproductives through cuticular hydrocarbons in the ant Odontomachus brunneus

Eusociality is characterized by a reproductive division of labor, wherein workers respond to the presence of reproductive individuals by refraining from reproduction themselves and restricting the reproductive efforts of others. Our understanding of how eusociality is maintained therefore depends on characterizing the mechanism by which workers detect the presence of a reproductive. Variations in cuticular hydrocarbons correspond to changes in reproductive ability in ants, and experimental studies are beginning to reveal the function of hydrocarbons as signals. In this study, we compare the cuticular hydrocarbon profiles of dominant and reproductive workers and queens of the ant Odontomachus brunneus with profiles of non-reproductive workers. Using split/reunification tests we document the existence of worker policing in both queenless and queenright colonies; supernumerary reproductives were treated aggressively by nestmates. Finally, we induce aggression and replicate queen-like submissive nestmate responses by supplementing the hydrocarbon profile of workers with (Z)-9-nonacosene, a compound that was significantly more abundant on the cuticles of reproductives. In three bioassays, we compare this manipulation to various control manipulations of the hydrocarbon profile and demonstrate that workers gauge the reproductive activity of nestmates through changes in their cuticular hydrocarbon profiles.     

Worker size-related task partitioning in the foraging strategy of a seed-harvesting ant species

Messor bouvieri is a seed-harvesting ant species in which workers forage in trails from the nest to a search area. A previous observation of seed transfer events between workers returning to the nest suggested potential task partitioning. In this study, we describe seed transportation and analyze the role of task partitioning in the foraging strategy of this species in terms of seed intake efficiency in relation to costs and benefits based on transport speed and task reliability. We assess the harvesting efficiency of task partitioning by comparing cooperative seed transport (CST) and individual seed transport (IST) events. Our results show task partitioning in the form of a sequence of transfer events among workers going from the search area to the nest. Importantly, and despite the weak worker polymorphism of this species, this sequence involved workers of different sizes, with seeds usually being passed along from smaller to larger workers. In addition, we show that small workers are better at finding seeds (spend less time finding a seed), and large workers are better at transporting them (were faster when walking back to the nest and lost fewer seeds). However, we failed to demonstrate that workers of different sizes are specialized in performing the task in which they excel. Overall, sequential CST in M. bouvieri results in a greater seed intake because seed search time decreases and task reliability increases, compared to IST. The determinants and adaptive benefits of CST are discussed.     

Worker discrimination among queens in newly founded colonies of the fire ant Solenopsis invicta

Although colonies of the fire ant Solenopsis invicta are often founded by small groups of queens, all but one of the queens are soon eliminated due to worker attacks and queen fighting. The elimination of supernumerary queens provides an important context for tests of discrimination by the workers, since the outcome of these interactions strongly affects the workers' inclusive fitness. To test whether workers in newly founded colonies discriminate among nestmate queens, paired cofoundresses were narrowly separated by metal screens that prevented direct fighting, but through which the workers could easily pass. Soon after the first workers completed development, they often attacked one of the queens; these attacks were strongly associated with queen mortality. When one queen's brood was discarded, so that the adult workers were all the daughters of just one queen, the workers were significantly less likely to bite their mother than the unrelated queen; however, this tendency was comparatively weak. Queens kept temporarily at a higher temperature to increase their rate of investment in brood-rearing lost weight more rapidly than paired queens and were subsequently more likely to be attacked and killed by workers. Workers were more likely to bite queens that had been temporarily isolated than queens that remained close to brood and workers. When queens were not separated by screens, the presence of workers stimulated queen fights. These results show that workers discriminate strongly among equally familiar queens and that discrimination is based more on the queens' condition and recent social environment than on kinship. Received: 9 June 1998 / Accepted after revision: 10 October 1998     

Orientation and recruitment behavior in the ponerine ant Pachycondyla tesserinoda (Emery): laying of individual-specific trails during tandem running

Summary Tandem leaders of Pachycondyla tesserinoda mark their way during nest moving. While scouting and foraging for food in an unknown area, chemical orientation is used as well. The origin of the trail substance could not be identified. Secretions of gastral exocrine glands did not induce trail following behavior. Nor do these secretions elicit tandem-following reactions; rather the latter is released by the general body surface odor. The trail substance and the substance used for marking the nest entrance are colony-specific. Moreover, individual tandem leaders recognize and show a preference for their own trails. This extraordinary effect is independent of the age of the trail. P. tesserinoda workers search individually for food and new nest sites. Targets which are important for the colony are directly shown to nestmates by tandem running. Due to this type of foraging and scouting individual-specific trails may be advantageous for this ant.     

A trail pheromone component of the African stink ant, Pachycondyla (Paltothyreus) tarsata Fabricius (Hymenoptera: Formicidae: Ponerinae)

Summary. The African stink ants (Pachycondyla tarsata) lay recruitment trails with secretions from sternal glands. The glandular secretions consist of 10 compounds, 9 of which have been chemically identified. One of the substances, 9-heptadecanone, elicits trail following behavior in P. tarsata workers that have before been stimulated by a sucessful scout ant. Received 7 August 1998; accepted 24 November 1998.     

Worker reproduction in the ponerine ant Ophthalmopone berthoudi: an alternative form of eusocial organization

Summary Colonies of the ponerine Ophthalmopone berthoudi were collected throughout the year. The queen caste is absent. Dissection of large numbers of workers revealed that many of them (up to 100 in one nest) are inseminated and produce eggs. The ovaries are small and contain very few mature oocytes, indicating that there is a slow rate of egg-laying. Workers are produced throughout the year, and all are capable of becoming functional reproductives. However, only those that are sexually-attractive during the limited period of male activity become mated. Thus the percentages of mated laying workers (=gamergates) fluctuate seasonally (Fig. 2). Successive generations of gamergates do not overlap. Observation of nests in the field and in the laboratory indicated that gamergates were never active above ground. There is no aggression between them, and their numbers are not socially regulated. There are few interactions between gamergates and non-reproductive workers, and the former do not acquire more food during termite meals. The notion of parental oppression is undermined by the complete loss of the meen caste, while the nature of the breeding system of this ant leads to the prediction of low relatedness between nestmates.     

Social rank, fecundity and lifetime reproductive success in wild European rabbits (Oryctolagus cuniculus)

Wild European rabbits of both sexes have separate linear rank orders, which are established and maintained by intensive fights. This paper presents data from a 14-year study (1987-2000) on the population physiology and behavioural ecology of wild rabbits living in a 22,000 m² enclosure and focuses on the relationship between social rank and reproduction in females. Group composition, social ranks, fecundity and reproductive success were known for all females (n=197) from the outset of their first reproductive season at an age of about 300 days until their death. The annual reproductive success of females was influenced to a large extent by their social rank. This depended on two effects of about equal strength: a higher fecundity of high-ranking females and a lower mortality of their offspring between birth and adulthood. The lifetime reproductive success of the females varied greatly. Only about 50% of all females had any reproductive success (range: on to nine adult offspring). The social rank achieved by the females during their first reproductive season substantially influenced their lifetime reproductive success: The mean reproductive lifespan and lifetime fitness of high-ranking females (ranks 1 and 2) was about 60% greater than that of lower-ranking females, although many of the latter also gained dominant positions in subsequent years.     

Changes in the cuticular hydrocarbons of incipient reproductives correlate with triggering of worker policing in the bulldog ant Myrmecia gulosa

In social insects, conflicts over male parentage can be resolved by worker policing. However, the evolution of policing behavior is constrained by the ability of individuals to identify reproductive nestmates, or their eggs. We investigated the occurrence of worker policing and its underlying chemical communication in the bulldog ant Myrmecia gulosa. Although workers have functional ovaries and can lay male-destined eggs, they do not reproduce in queenright colonies. To determine if their sterility is a consequence of worker policing, we experimentally induced worker reproduction in the presence of a queen. Some individuals were seized and immobilized by nestmates, and sometimes killed as a consequence. Although the ovarian development of immobilized individuals was variable, their cuticular hydrocarbon profiles were intermediate between reproductive and nonreproductive workers, indicating they were in the process of starting to reproduce. Approximately 29% of these incipient reproductive workers were successfully policed. To test for policing on eggs, we transferred viable worker eggs to queenright colonies and monitored their acceptance. Furthermore, we compared the surface hydrocarbons of the different types of eggs to determine whether these chemicals could be involved in egg recognition. We found that although there were differences in hydrocarbon profiles and discrimination between queen and worker-laid eggs, viable eggs were not destroyed. Our results strongly support the idea that cuticular hydrocarbons are involved in the policing of reproductive workers. A low level of worker policing appears sufficient to select for self-restraint in workers when few fitness benefits are gained by selfish reproduction. Policing of eggs may thus be unnecessary.