Queen–worker caste ratio depends on colony size in the pharaoh ant (<Emphasis Type="Italic">Monomorium pharaonis</Emphasis>)

The success of an ant colony depends on the simultaneous presence of reproducing queens and non-reproducing workers in a ratio that will maximize colony growth and reproduction. Despite its presumably crucial role, queen–worker caste ratios (the ratio of adult queens to workers) and the factors affecting this variable remain scarcely studied. Maintaining polygynous pharaoh ant (Monomorium pharaonis) colonies in the laboratory has provided us with the opportunity to experimentally manipulate colony size, one of the key factors that can be expected to affect colony level queen–worker caste ratios and body size of eclosing workers, gynes and males. We found that smaller colonies produced more new queens relative to workers, and that these queens and workers both tended to be larger. However, colony size had no effect on the size of males or on the sex ratio of the individuals reared. Furthermore, for the first time in a social insect, we confirmed the general life history prediction by Smith and Fretwell (Am Nat 108:499–506, 1974) that offspring number varies more than offspring size. Our findings document a high level of plasticity in energy allocation toward female castes and suggest that polygynous species with budding colonies may adaptively adjust caste ratios to ensure rapid growth.     

Behavioral and Chemical Correlates of Long-Term Queen Adoption in the Facultative Polygynous Ant Ectatomma tuberculatum

In ants, queen adoption is a common way of achieving secondary polygyny but the mechanisms involved are little known. Here we studied the process of long-term adoptions of alien queens in the facultative polygynous ant Ectatomma tuberculatum. In eight out of 10 successful adoption experiments, all the introduced queens showed similar behavior and fecundity as the resident queens even after 2 months, indicating complete integration into the colony. Chemical analysis revealed that the cuticular hydrocarbon profiles of resident and introduced queens were clearly distinct from those of workers and that they did not change after adoption. We propose that queen-specific cuticular hydrocarbon profile may represent a reliable signal of queen’s fertility and discuss about the evolution of high level of queen acceptance in E. tuberculatum.     

Development rate and brood production in haplo- and pleometrotic colonies of <Emphasis Type="Italic">Oecophylla smaragdina</Emphasis>

Pleometrosis (colony founding by multiple queens) may improve life history characteristics that are important for early colony survival. When queens unite their initial brood, the number of workers present when incipient colonies open may be higher than for single queen colonies. Further, the time until the first worker emerges may shorten. For territorial species and species that rob brood from neighbouring colonies, a faster production of more workers may improve the chance of surviving intraspecific competition. In this study, the time from the nuptial flight to the emergence of the first worker in incipient Oecophylla smaragdina Fabr. colonies founded by 1–5 queens was compared and the production of brood during the first 68 days after the nuptial flight was assessed. Compared to haplometrotic colonies, pleometrotic colonies produced 3.2 times more workers, their first worker emerged on average 4.3 days (8%) earlier and the queen’s per capita egg production almost doubled. Further, colony production was positively, correlated with the number of founding queens and time to worker emergence was negatively correlated. These results indicate that pleometrotic O. smaragdina colo-nies are competitively superior to haplometrotic colonies as they produce more workers faster and shorten the claustral phase, leading to increased queen fecundity.     

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.     

Bigger Helpers in the Ant Cataglyphis bombycina: Increased Worker Polymorphism or Novel Soldier Caste?

Introduction

The mechanisms by which development favors or constrains the evolution of new phenotypes are incompletely understood. Polyphenic species may benefit from developmental plasticity not only regarding ecological advantages, but also potential for evolutionary diversification. For instance, the repeated evolution of novel castes in ants may have been facilitated by the existence of alternative queen and worker castes and their respective developmental programs.

Material and Methods

Cataglyphis bombycina is exceptional in its genus because winged queens and size-polymorphic workers occur together with bigger individuals having saber-shaped mandibles. We measured seven body parts in more than 150 individuals to perform a morphometric analysis and assess the developmental origin of this novel phenotype.

Results

Adults with saber-shaped mandibles differ from both workers and queens regarding the size of most body parts. Their relative growth rates are identical to workers for some pairs of body parts, and identical to queens for other pairs of body parts; critical sizes differ in all cases.

Conclusions

Big individuals are a third caste, i.e. soldiers, not major workers. Novel traits such as elongated mandibles are combined with a mix of queen and worker growth rates. We also reveal the existence of a dimorphism in the queen caste (microgynes and macrogynes). We discuss how novel phenotypes can evolve more readily in the context of an existing polyphenism. Both morphological traits and growth rules from existing queen and worker castes can be recombined, hence mosaic phenotypes are more likely to be viable. In C. bombycina, such a mosaic phenotype appears to function both for defense (saber-shaped mandibles) and fat storage (big abdomen). Recycling of developmental programs may have contributed to the morphological diversity and ecological success of ants.     

Aggregation and adult transportation in disturbed colonies of <Emphasis Type="Italic">Wasmannia auropunctata</Emphasis> Roger (Hymenoptera,Formicidae)

Disturbance of W. auropunctata colonies first produced, as expected, significant dispersal of workers and queens, but this was soon followed by a radiate aggregation of workers, heads to center, in groups of 5 – 8 individuals, all with mandibles widely open. Queens did not aggregate, but instead searched for the grouped workers, antennating the gaster of a selected individual; after this, workers began to slowly board onto the queen’s body. This behavior was repeated with other groups, until the queen could not bear any more workers. This behavior is registered for the first time for adult ants, and is interpreted as a unique strategy to reorganize the nest or to establish a new colony. The open mandibles may indicate the release of alkylpyrazine, a known aggregation pheromone. Received 8 January 2007; revised 28 February 2007; accepted 2 March 2007.     

Mating system evolution and worker caste diversity in Pheidole ants

The efficiency of social groups is generally optimized by a division of labour, achieved through behavioural or morphological diversity of members. In social insects, colonies may increase the morphological diversity of workers by recruiting standing genetic variance for size and shape via multiply mated queens (polyandry) or multiple‐breeding queens (polygyny). However, greater worker diversity in multi‐lineage species may also have evolved due to mutual worker policing if there is worker reproduction. Such policing reduces the pressure on workers to maintain reproductive morphologies, allowing the evolution of greater developmental plasticity and the maintenance of more genetic variance for worker size and shape in populations. Pheidole ants vary greatly in the diversity of worker castes. Also, their workers lack ovaries and are thus invariably sterile regardless of the queen mating frequency and numbers of queens per colony. This allowed us to perform an across‐species study examining the genetic effects of recruiting more patrilines on the developmental diversity of workers in the absence of confounding effects from worker policing. Using highly variable microsatellite markers, we found that the effective mating frequency of the soldier‐polymorphic P. rhea (avg. me_N = 2.65) was significantly higher than that of the dimorphic P. spadonia (avg. me_N = 1.06), despite a significant paternity skew in P. rhea (avg. B = 0.10). Our findings support the idea that mating strategies of queens may co‐evolve with selection to increase the diversity of workers. We also detected patriline bias in the production of different worker sizes, which provides direct evidence for a genetic component to worker polymorphism.     

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.     

Spatio-temporal variation of colony structure and eusociality level of the Japanese sweat bee<Emphasis Type="Italic"> Lasioglossum</Emphasis> (<Emphasis Type="Italic">Evylaeus</Emphasis>)<Emphasis Type="Italic"> duplex</Emphasis> (Hymenoptera: Halictidae)

Colony structure and eusociality level of the sweat bee Lasioglossum (Evylaeus) duplex were studied in 2001 in Sapporo and Assabu, Hokkaido, northern Japan. Sakagami and his colleagues had also studied this species in Sapporo in 1957–1968. Brood size, sex ratio and queen–worker size dimorphism were geographically and temporally variable, indicating spatio-temporal variation at the eusociality level. Inseminated workers constituted only 7.9% of the populations in 1957–1968 Sapporo but about 60% in 2001 in Sapporo and Assabu. A few of the inseminated workers were believed to leave natal nests for independent colony founding. Thus, partial bivoltinism is likely in this sweat bee species. The presence of workers with developed ovaries and/or corpora lutea suggests the occurrence of worker oviposition.     

Nesting patterns,ecological correlates of polygyny and social organization in the neotropical arboreal ant <Emphasis Type="Italic">Odontomachus hastatus</Emphasis> (Formicidae,Ponerinae)

Queen number varies in the population of O. hastatus in SE Brazil. Here, we evaluate how nesting ecology and colony structure are associated in this species, and investigate how reproduction is shared among nestmate queens. Queen number per colony is positively correlated with nesting space (root cluster of epiphytic bromeliads), and larger nest sites host larger ant colonies. Plant samplings revealed that suitable nest sites are limited and that nesting space at ant-occupied bromeliads differs in size and height from the general bromeliad community. Dissections revealed that queens in polygynous colonies are inseminated, have developed ovaries, and produce eggs. Behavioral observations showed that reproduction in polygynous colonies is mediated by queen–queen agonistic interactions that include egg cannibalism. Dominant queens usually produced more eggs. Field observations indicate that colonies can be initiated through haplometrosis. Polygyny in O. hastatus may result either from groups of cofounding queens (pleometrosis) or from adoption of newly mated queens by established colonies (secondary polygyny). Clumping of bromeliads increases nest space and probably adds stability through a strong root system, which may promote microhabitat selection by queens and favor pleometrosis. Rainstorms that frequently knock down bromeliads can be a source of colony break-up and may promote polygyny. Bromeliads are limited nest sites and may represent a risk for young queens leaving a suitable nest, thus favoring secondary polygyny. Although proximate mechanisms mediating queen number are poorly understood, this study suggests that heterogeneous microhabitat conditions probably contribute to the coexistence of variable forms of social structure in O. hastatus.     

Notes on the biology of the Oriental amblyoponine ant Myopopone castanea: Queen‐worker dimorphism,worker polymorphism and larval hemolymph feeding by workers (Hymenoptera: Formicidae)

Queen‐worker dimorphism, worker polymorphism and worker behavior in the Oriental amblyoponine ant Myopopone castanea were studied. Queen body size was large (head width, 3.0 mm) with 24 to 32 ovarioles while workers showed a remarkable size variation in both body size (head width 1.48 to 2.18 mm) and ovariole number (6 to 22). Both head width and abdomen width showed allometric growth against thorax width. Workers performed larval hemolymph feeding as was described for Amblyopone silvestrii queens.     

A newly-discovered mode of colony founding among fire ants

Summary Queen ants start new colonies either unassisted by workers (independent founding), assisted by workers from their natal nest (dependent founding), or assisted by the workers of other species (dependent, socially parasitic). The monogyne form of the fire ant,Solenopsis invicta, founds independently in summer, but in the fall it also produces a few sexuals some of which overwinter, then fly and mate in early spring. These overwintered queens lack the nutritional reserves and behaviors for independent colony founding. Rather, they seek out unrelated, mature, orphaned colonies, enter them and exploit the worker force to found their own colony through intraspecific social parasitism. Success in entering orphaned colonies is higher when these lack overwintered female alates of their own. When such alates are present, orphaning causes some to dealate and become uninseminated replacement queens, usually preventing entry of unrelated, inseminated replacement queens. Such colonies produce large, all-male broods. Successful entry of a parasitic queen robs the host colony of this last chance at reproductive success. Only overwintered sexuals take part in this mode of founding.     

Development and evolution of caste dimorphism in honeybees – a modeling approach

The difference in phenotypes of queens and workers is a hallmark of the highly eusocial insects. The caste dimorphism is often described as a switch‐controlled polyphenism, in which environmental conditions decide an individual's caste. Using theoretical modeling and empirical data from honeybees, we show that there is no discrete larval developmental switch. Instead, a combination of larval developmental plasticity and nurse worker feeding behavior make up a colony‐level social and physiological system that regulates development and produces the caste dimorphism. Discrete queen and worker phenotypes are the result of discrete feeding regimes imposed by nurses, whereas a range of experimental feeding regimes produces a continuous range of phenotypes. Worker ovariole numbers are reduced through feeding‐regime‐mediated reduction in juvenile hormone titers, involving reduced sugar in the larval food. Based on the mechanisms identified in our analysis, we propose a scenario of the evolutionary history of honeybee development and feeding regimes.     

Population and colony structure and morphometrics in the queen dimorphic harvester ant, <Emphasis Type="Italic">Pogonomyrmex pima</Emphasis>

The North American seed-harvester ant Pogonomyrmex (Ephebomyrmex) pima displays a dimorphism that consists of winged (alate) and wingless (intermorph) queens; both types of queens are fully reproductive. Microsatellite allele frequencies and a mitochondrial phylogeny demonstrate (1) alate and intermorph queens represent an intraspecific wing polymorphism, and (2) an absence of assortative mating and inbreeding by males. Surveys at our field site in southcentral Arizona, USA, demonstrated that only one type of queen (intermorph or dealate) occurred in each colony, including those excavated during the season in which reproductive sexuals were present. Colony structure appeared to vary by queen type as most intermorph colonies contained multiple mated queens. Alternatively, dealate queen colonies rarely contained a mated queen. Our inability to find mated dealate queens in these colonies probably resulted from difficulty in excavating the entire colony and reproductive queen, especially given that these colonies were only excavated over one day. A morphometric analysis demonstrated that intermorph queens are intermediate in size to that of workers and alate queens, but that intermorph queens retain all of the specialized anatomical features of alate queens (except for wings). Some colonies had queens that foraged and performed nest maintenance activities, and these queens sometimes accounted for a significant portion of colony foraging trips. Dissections revealed that these queens were uninseminated; some of these queens produced males in the laboratory. Received 24 October 2006; revised 1 December 2006; accepted 8 December 2006.     

Colony genetic structure in the Australian jumper ant <Emphasis Type="Italic">Myrmecia pilosula</Emphasis>

Eusocial insects vary significantly in colony queen number and mating frequency, resulting in a wide range of social structures. Detailed studies of colony genetic structure are essential to elucidate how various factors affect the relatedness and the sociogenetic organization of colonies. In this study, we investigated the colony structure of the Australian jumper ant Myrmecia pilosula using polymorphic microsatellite markers. Nestmate queens within polygynous colonies, and queens and their mates, were generally unrelated. The number of queens per colony ranged from 1 to 4. Queens were estimated to mate with 1–9 inferred and 1.0–11.4 effective mates. This is the first time that the rare co-occurrence of polygyny and high polyandry has been found in the M. pilosula species group. Significant maternity and paternity skews were detected at the population level. We also found an isolation-by-distance pattern, and together with the occurrence of polygynous polydomy, this suggests the occurrence of dependent colony foundation in M. pilosula; however, independent colony foundation may co-occur since queens of this species have fully developed wings and can fly. There is no support for the predicted negative association between polygyny and polyandry in ants.     

Habitat structure,dispersal strategies and queen number in two boreal Leptothorax ants

In two nearctic ants, Leptothorax canadensis and Leptothorax sp. A, young queens may either found their own nest solitarily after mating or seek adoption into an established colony. Whether a queen disperses or not is associated with genetically determined queen morphology in Leptothorax sp. A. Whereas a majority of winged queens attempt solitary colony founding after mating, most wingless, intermorphic queens return to their maternal nests and new colonies are founded by budding after hibernation. The latter strategy appears to be correlated with patchy, isolated habitats, whereas in extended boreal forests dispersal on the wing is probably more common. Alternative dispersal strategies strongly affect the average number of queens per colony and seasonal fluctuations of colony structure.     

COMPLEX COLONY STRUCTURE IN SOCIAL INSECTS: I. ECOLOGICAL DETERMINANTS AND GENETIC CONSEQUENCES

For social insect species, intraspecific variation in colony social structure provides an opportunity to relate the evolution of social behavior to ecological factors. The species Myrmica punctiventris is a cavity-dwelling forest ant that exhibits very different colony structures in two populations in the northeastern United States. Combined data from seasonal censuses, allozyme electrophoresis, and worker hostility tests showed that a population of M. punctiventris in Vermont was strictly monogynous and seasonally polydomous. The same procedures showed that a population of M. punctiventris in New York was facultatively polygynous and predominantly monodomous. Genetic relatedness among colony-mates was not different from Hamilton's expected values in the Vermont population and was consistent with little exchange of ants between colonies and single-mating of queens. In contrast, relatedness was lower in New York, and examination of nest-mate genotypes revealed exchange of ants between colonies, high rates of colony loss and replacement of queens, or multiple-mating of queens. The genetic structure of the Vermont population was consistent with no inbreeding, but in New York, the population genetic structure reflected microgeographic subdivision and inbreeding. Previous study of the ant communities at these sites implicates nest-site limitation in New York as a primary constraint on social structure.     

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.     

Gene flow is maintained by polyandry and male dispersal in the army ant <Emphasis Type="Italic">Eciton burchellii</Emphasis>

The combination of haplodiploidy, complementary sex determination and eusociality constrains the effective population size (N _e) of social Hymenoptera far more than in any other insect group. Additional limitations on N _e occur in army ants since they have wingless queens and colony fission, both of which are factors causing restricted maternal gene flow and high population viscosity. Therefore, winged army ant males gain a particular significance to ensure dispersal, facilitate gene flow and avoid inbreeding. Based on population genetic analyses with microsatellite markers, we studied a population of the Neotropical army ant Eciton burchellii, finding a high level of heterozygosity, weak population differentiation and no evidence for inbreeding. Moreover, by using sibship reconstruction analyses, we quantified the actual number of male contributing colonies represented in a queen’s mate sample, demonstrating that, through extreme multiple mating, the queens are able to sample the genes of males from up to ten different colonies, usually located within an approximate radius of 1 km. We finally correlated the individual mating success of each male contributing colony with the relative siring success of individual males and found a significant colony-dependent male fitness component. Our results imply that the dispersal and mating system of these army ants seem to enhance gene flow and minimise the deleterious effects associated with small effective population size. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.