Colony fusion and worker reproduction after queen loss in army ants

Theory predicts that altruism is only evolutionarily stable if it is preferentially directed towards relatives, so that any such behaviour towards seemingly unrelated individuals requires scrutiny. Queenless army ant colonies, which have anecdotally been reported to fuse with queenright foreign colonies, are such an enigmatic case. Here we combine experimental queen removal with population genetics and cuticular chemistry analyses to show that colonies of the African army ant Dorylus molestus frequently merge with neighbouring colonies after queen loss. Merging colonies often have no direct co-ancestry, but are on average probably distantly related because of overall population viscosity. The alternative of male production by orphaned workers appears to be so inefficient that residual inclusive fitness of orphaned workers might be maximized by indiscriminately merging with neighbouring colonies to increase their reproductive success. We show that worker chemical recognition profiles remain similar after queen loss, but rapidly change into a mixed colony Gestalt odour after fusion, consistent with indiscriminate acceptance of alien workers that are no longer aggressive. We hypothesize that colony fusion after queen loss might be more widespread, especially in spatially structured populations of social insects where worker reproduction is not profitable.     

Visual prey discrimination of queen and worker ants by a generalist lizard

Laboratory experiments were conducted to test the hypothesis that widelyforaging generalist lizard,Eumeces okadae, visually discriminates palatable queen ants from unpalatable worker ants. The workers ofCamponotus japonicus andLasius niger were rejected on sight, while the queens of both species were eaten with little prior chemical examination by tongue flicks or licks. Comparison of the lizards' responses towards the workers and wingless queens of similar size indicated that neither body size nor presence or absence of wings accounted for difference in responses toward the 2 ant castes. The lizards probably discriminated different ant castes by body proportions.     

Sex allocation in fungus‐growing ants: worker or queen control without symbiont‐induced female bias

The fungal cultivars of fungus‐growing ants are vertically transmitted by queens but not males. Selection would therefore favor cultivars that bias the ants’ sex ratio towards gynes, beyond the gyne bias that is optimal for workers and queens. We measured sex allocation in 190 colonies of six sympatric fungus‐growing ant species. As predicted from relatedness, female bias was greater in four singly mated Sericomyrmex and Trachymyrmex species than in two multiply mated Acromyrmex species. Colonies tended to raise mainly a single sex, which could be partly explained by variation in queen number, colony fecundity, and fungal garden volume for Acromyrmex and Sericomyrmex, but not for Trachymyrmex. Year of collection, worker number and mating frequency of Acromyrmex queens did not affect the colony sex ratios. We used a novel sensitivity analysis to compare the population sex allocation ratios with the theoretical queen and worker optima for a range of values of k, the correction factor for sex differences in metabolic rate and fat content. The results were consistent with either worker or queen control, but never with fungal control for any realistic value of k. We conclude that the fungal symbiont does not distort the ants’ sex ratio in these species.     

Environmental and genetic influences on queen and worker body size in the social wasp Vespula maculifrons

Many social insects exhibit morphologically distinct worker and queen castes that perform different functions. These functional differences may generate unique selection regimes operating on body size. For example, queens may be under directional selection for large body size, whereas directional selection on worker body size may be limited. Such contrasting selection pressures may differentially affect levels of genetic variation associated with size variation in the two castes. This study sought to determine if genetic effects underlying phenotypic differences varied between the worker and queen castes of the social wasp Vespula maculifrons. We predicted that directional selection would remove genetic variation associated with size differences in the queen caste, whereas a lack of directional selection would tend to maintain genetic variation associated with size differences in the worker caste. We thus (1) calculated broad and narrow sense heritabilities for several morphological traits, (2) examined whether some paternal genotypes produced more morphologically diverse offspring than others, and (3) determined whether trait size variation was associated with genetic variation within colonies. We found that few morphological traits were significantly heritable, indicating that little genetic variance for those traits existed within our study population. We also found that some patrilines produced more morphologically variable offspring than others, suggesting a role of genotype in phenotypic plasticity. And finally, no significant correlations between genetic diversity arising from multiple mating by queens within colonies and trait variation in either caste were found. Overall, our findings indicate a weak effect of genotype on both worker and queen body size variation and are suggestive of a large environmental influence on morphological trait size. Moreover, our results do not indicate that levels of genetic variation underlying size variation differ substantially between castes in this species.     

Colony genetic structure and queen mating frequency in fire ants of the subgenus Solenopsis (Hymenoptera: Formicidae)

Colony genetic structure was studied in natural populations of three fire ant taxa, Solenopsis richteri Forel, S. geminata (Fabr.), and hybrid S. invicta/richteri , using allozyme markers. All colonies studied exhibited arrays of female genotypes predicted under a model of monogyny (single functional queen) and monoandry (single insemination of queens). Males produced in the colonies appear to originate exclusively from the foundress queen, rather than from any virgin females present in the colonies. Thus these social insect colonies represent simple, albeit enormous, family groups. Single insemination and foundress parentage of males appear to be conserved reproductive traits in the subgenus Solenopsis , whereas another major determinant of colony genetic structure, the number of functional queens, is evolutionarily labile in this group.     

Caste control through worker attack in the antMyrmica

Summary Simulation of the way workers attack big larvae in spring has produced evidence that the mechanical component of this alone is sufficient to induce earlier metamorphosis with a consequent switch from queen to worker formation.As the jaws are used and the mandibular glands contain a proteinase normally used in the external digestion of prey, it is highly probable that this, too, is involved in attack. It is shown that the mandibular gland material causes major damage to the cuticle of the larva and may cause it to lose water.Like queens, workers contain fatty acids in their heads and the chain length frequency is quite similar. In both castes the commonest is oleic acid. Applied topically this enters the larva, reduces its growth rate and damages its water-proofing. Water-impermeability can be restored by workers.These fatty acids, however, are not very abundant inMyrmica. They are present in the pharyngeal gland which is used to feed larvae and is very palatable. There is no evidence that it is used topically as a poison though it is certainly able to pass straight in through the larval cuticle; if implanted it is harmless.The mandibular glands of queens and workers have been removed surgically. Workers are still able to rear larvae quite well and are susceptible to the influence of queens, but to a lesser degree than normal. This is either because they are undernourished, as the gland is their main oral source of proteinase, or because they need the mandibular chemicals to raise the intensity of their group aggression.the pharyngeal glands of queens and workers have also been removed. This destroys much of the worker drive towards larvae and leads to their gradual emaciation, probably because some dligestion and absorption takes place there. Larvae lose weight, too, and so the influence of queens on broodrearing cannot be tested.

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Résumé En simulant la manière dont les ouvrières attaquent au printemps les grandes larves, on montre que les stimulations mécaniques seules suffisent à induire prématurément la métamorphose avec production d'ouvrières ou bien de reine.Puisque les mandibules sont utilisées et que les glandes mandibulaires contiennent une protéinase, utilisée normalement pour la digestion externe des proies, il est hautement probable que cela, aussi, intervienne dans l'attaque. Il est montré que la subtance des glandes mandibulaires occasionne les principaux dégâts à la cuticule de la larve et peut induire des pertes d'eau.Comme celle des reines, la tête des ouvrières contient des acides gras et leur répartition en fonction de la longueur des chaînes est rigoureusement identique. Dans les deux castes, le plus commun est l'acide oléique. Appliqué localement, celui-ci pénètre dans la larve, réduit son taux de croissance et perturbe son imperméabilité à l'eau. L'imperméabilité à l'eau peut être rétablie par les ouvrières.Cependant, ces acides gras ne sont pas très abondants chezMyrmica. Ils sont présents dans la glande pharyngienne qui est utilisée dans le nourrissement des larves et est d'un goût agréable. Il n'y a aucune preuve qu'elle soit utilisée localement comme poison, bien qu'elle soit certainement capable de traverser directement la cuticule larvaire; implantée, elle est inoffensive.L'ablation chirurgicale des glandes mandibulaires de reines et d'ouvrières a été faite. Les ouvrières sont encore tout à fait capables d'élever des larves et sont sensibles à l'influence des reines, mais à un degré moindre que normal. Cela est dû soit à une sous-alimentation, car la glande est leur principale source orale de protéinase, soit parce qu'elles ont besoin des substances mandibulaires pour augmenter l'intensité de l'agression de leur groupe.Les glandes pharyngiennes de reines et d'ouvrières ont aussi, été enlevées. Cela annihile beaucoup de l'attirance des ouvrières vers les larves et mène à leur déssèchement progressif probablement parce qu'il y a alors une certaine digestion et absorption. Les larves perdent aussi du poids, et ainsi l'influence des reines sur l'élevage du couvain ne peut pas être étudiée.

     

Male reproductive investment and queen mating-frequency in fungus-growing ants

Sperm number and male accessory gland compounds are often importantdeterminants of male mating success but have been little studiedin social insects. This is because mating in social insectsis often difficult to manipulate experimentally, and first evidencefor an explicit influence of accessory gland secretions on malemating success in social insects was obtained only recently.Here we perform a comparative analysis of male sexual organsacross 11 species of attine fungus-growing ants, representingboth genera with single- and multiple-queen mating. We foundthat the general morphology of the male sexual organs was verysimilar across all species, but the relative sizes of the accessoryglands and the sperm-containing accessory testes vary significantlyacross species. Small testes and large accessory glands characterizespecies with singly mated queens, whereas the opposite is foundin species with multiply mated queens. However, in the socialparasite Acromyrmex insinuator, in which queens have secondarilyreverted to single mating, males have accessory gland characteristicsreminiscent of the lower attine ants, but without having significantlyreduced their investment in sperm production. We hypothesizethat the main function of accessory gland compounds in attineants is to monopolize male paternity in similar ways as knownfrom other social insects. This would imply that the evolutionof polyandry in the terminal clade of the fungus-growing ants(the leafcutter ants) has resulted in selection for decreasedinvestment by males in accessory gland secretions and increasedinvestment in sperm number, in response to sperm competitionfor sperm storage.     

Alternative reproductive strategies: a queen perspective in ants

Ant colonies are commonly thought to have a stable and simple family structure, with one or a few egg-laying queens and their worker daughters. However, recent genetic studies reveal that the identity of breeding queens can vary over time within colonies. In several species, some queens are apparently specialized to enter established colonies instead of initiating a new colony on their own. The previously overlooked occurrence of queen turnover within colonies has important consequences not only on the genetic structure and nature of kin conflict within colonies, but also on the evolution of social parasitism.     

Sex allocation conflict in ants: when the queen rules

Insect societies are paramount examples of cooperation, yet they also harbor internal conflicts whose resolution depends on the power of the opponents. The male-haploid, female-diploid sex-determining system of ants causes workers to be more related to sisters than to brothers, whereas queens are equally related to daughters and sons. Workers should thus allocate more resources to females than to males, while queens should favor an equal investment in each sex. Female-biased sex allocation and manipulation of the sex ratio during brood development suggest that workers prevail in many ant species. Here, we show that queens of Formica selysi strongly influenced colony sex allocation by biasing the sex ratio of their eggs. Most colonies specialized in the production of a single sex. Queens in female-specialist colonies laid a high proportion of diploid eggs, whereas queens in male-specialist colonies laid almost exclusively haploid eggs, which constrains worker manipulation. However, the change in sex ratio between the egg and pupae stages suggests that workers eliminated some male brood, and the population sex-investment ratio was between the queens' and workers' equilibria. Altogether, these data provide evidence for an ongoing conflict between queens and workers, with a prominent influence of queens as a result of their control of egg sex ratio.     

Effect of honey bee queen mating condition on worker ovary activation

The presence of the honey bee queen reduces worker ovary activation. When the queen is healthy and fecund, this is interpreted as an adaptive response as workers can gain fitness from helping the queen raise additional offspring, their sisters. However, when the queen is absent, workers activate their ovaries and lay unfertilized eggs that become males. Queen pheromones are recognised as a factor affecting worker ovary activation. Recent work has shown that queen mandibular pheromone composition changes with queen mating condition and workers show different behavioural responses to pheromone extracts from these queens. Here, we tested whether workers reared in colonies with queens of different mating condition varied in level of ovary activation. We also examined the changes in the chemical composition of the queen mandibular glands to determine if the pheromone blend varied among the queens. We found that the workers activated their ovaries when queens were unmated and had lower ovary activation when raised with mated queens, suggesting that workers detect and respond adaptively to queens of differing mating status. Moreover, variation in queen mandibular gland’s chemical composition correlated with the levels of worker ovary activation. Although correlative, this evidence suggests that queen pheromone may act as a signal of queen mating condition for workers, in response to which they alter their level of ovary activation.     

The importance of workers for queen hibernation survival in Camponotus ants

The higher proportion of polygynous ant species in northern areas indicates that cold climates influence queen number per colony. It is unclear, however, what ecological and physiological factors facilitate the dominance of polygynous species in cold climates. This is the case in two common arboreal ants in Japan-Camponotus yamaokai and C. nawai-which are quite similar in morphology, but different in social structure and geographical distribution. Polygynous C. yamaokai inhabits colder areas, whereas monogynyous C. nawai inhabits warmer climates. We compared queen survival in both ants at low temperature to evaluate whether interspecific difference in cold tolerance can explain the geographical distribution. We examined the influence of cohabitation with other individuals, as well as individual cold tolerance. Experimental groups with different caste compositions were prepared and maintained under conditions simulating in the laboratory climates of the northern limit of C. nawai. Wintering experiments revealed that C. yamaokai queens survived longer than C. nawai queens under solitary conditions, although half of the queens died in less than a month, even in C. yamaokai. Queens hibernating with workers survived longer than solitary queens, but queen number did not affect queen survival. Cohabitation with workers allowed 80% of C. yamaokai queens to survive more than two months. Under field conditions, monogynous C. nawai foundresses overwinter without workers, whereas new queens of polygynous C. yamaokai always overwinter with many workers. Thus, the geographical distribution of these ants appears to depend on the overwintering behavior of new queens.     

Influence of queen weight and colony origin on worker response in Solenopsis geminata

Abstract. The influence of weight and colony origin of the queen of Solenopsis geminata (F.) (Hymenoptera: Formicidae) on worker attraction is studied under laboratory conditions. In the first experiment, worker response to individual queens of different weight from the same colony is evaluated. Heavier queens are more attractive than smaller queens to their own workers. In subsequent experiments, the colony origin effect is investigated and worker response to a pair of queens of the same weight from the same or different colonies is compared. When queens are from the same colony, workers do not show a significant preference between queens. However, when queens are from a different colony, workers are significantly more attracted to their own queen than to the foreign queen. Finally, the response of workers to queens of different weight from the same or different colonies is investigated. In both cases, workers are significantly more attracted to a heavier queen than a lighter queen, even if the lighter queen is their own queen. A putative pheromonal component (E)‐6‐(1‐pentenyl)‐2H‐2‐pyranone, is not positively correlated with queen weight.     

cGMP modulates responses to queen mandibular pheromone in worker honey bees

Responses to social cues, such as pheromones, can be modified by genotype, physiology, or environmental context. Honey bee queens produce a pheromone (queen mandibular pheromone; QMP) which regulates aspects of worker bee behavior and physiology. Forager bees are less responsive to QMP than young bees engaged in brood care, suggesting that physiological changes associated with behavioral maturation modulate response to this pheromone. Since 3′,5′-cyclic guanosine monophosphate (cGMP) is a major regulator of behavioral maturation in workers, we examined its role in modulating worker responses to QMP. Treatment with a cGMP analog resulted in significant reductions in both behavioral and physiological responses to QMP in young caged workers. Treatment significantly reduced attraction to QMP and inhibited the QMP-mediated increase in vitellogenin RNA levels in the fat bodies of worker bees. Genome-wide analysis of brain gene expression patterns demonstrated that cGMP has a larger effect on expression levels than QMP, and that QMP has specific effects in the presence of cGMP, suggesting that some responses to QMP may be dependent on an individual bees’ physiological state. Our data suggest that cGMP-mediated processes play a role in modulating responses to QMP in honey bees at the behavioral, physiological, and molecular levels.     

Polyethism in social interactions in ants

Worker ants, which perform various tasks in their society (division of labour or polyethism), also demonstrate different types of antennal activity in trophallactic interactions. These differences concern the variability in successions of behavioural units performed by the worker in the course of substance transmission as well as in the sequential organization of these units. The antennal activity of an aged worker varies according to whether the ant gives food to another aged worker or to a callow worker. In the latter case, the donor's behaviour does not differ according to its social function (brood-tending worker or forager). A parallel can be established between these results and those obtained in a previous study of aggressive behaviour. This behaviour is liable to appear when a callow worker extracted from its society before hatching and therefore deprived of relations with aged workers of the society, is therefore placed with an aged worker. Polyethism is then manifested in social interactions, and not only in the division of labour between members of the society.     

Evolution of advanced social traits in phylogenetically basal ants: striking worker polymorphism and large queens in Amblyopone australis

The ecological success of ants results notably from their ability to derive several phenotypes from the same genotype. In higher ants, polyphenism can be extreme: high queen–worker size dimorphism and worker polymorphism allow for better success at colony founding by lone queens, better division of tasks, and ultimately access to new ecological niches. However in basal ants, polyphenism is much more limited, restricting them to narrow niches. It is often assumed that basal ants lack the ability to produce a pronounced polyphenism. In Amblyopone, most species nest and hunt centipedes in soil and leaf litter, and polyphenism is weak, e.g. A. pallipes. Here we studied A. australis, which forages and nests in rotten logs. Using analyses of morphometry and allometry, we showed that queen–worker dimorphism and worker polymorphism are much higher in A. australis than in A. pallipes. Workers of A. australis exhibit various sizes and shapes, and large individuals could be better adapted for digging galleries in rotten logs. Moreover, larger queens are probably more efficient during non-claustral colony foundation. We conclude that the evolution of advanced social traits typical of higher ants is also possible in basal ants, but it is not often selected for.     

Colony size predicts division of labour in attine ants

Division of labour is central to the ecological success of eusocial insects, yet the evolutionary factors driving increases in complexity in division of labour are little known. The size–complexity hypothesis proposes that, as larger colonies evolve, both non-reproductive and reproductive division of labour become more complex as workers and queens act to maximize inclusive fitness. Using a statistically robust phylogenetic comparative analysis of social and environmental traits of species within the ant tribe Attini, we show that colony size is positively related to both non-reproductive (worker size variation) and reproductive (queen–worker dimorphism) division of labour. The results also suggested that colony size acts on non-reproductive and reproductive division of labour in different ways. Environmental factors, including measures of variation in temperature and precipitation, had no significant effects on any division of labour measure or colony size. Overall, these results support the size–complexity hypothesis for the evolution of social complexity and division of labour in eusocial insects. Determining the evolutionary drivers of colony size may help contribute to our understanding of the evolution of social complexity.