Dynamics and Consequences of Hierarchy Formation in the Ant Leptothorax sp. A

In the Nearctic ant Leptothorax sp. A, aggressive interactions among wingless intermorphic queens and primarily winged gynomorphic queens lead to the formation of dominance hierarchies, in which the highest-ranking individual is the only egg-layer in a colony. Fighting occurs during two periods of the annual cycle: in late summer, newly adopted young queens are integrated into the colony's hierarchy; after hibernation, fighting resumes and the high aggressiveness of α-queens may now lead to the emigration of β and other middle-ranking queens. The α-position appears to be very stable over successive fighting periods, though an estimate of nestmate relatedness by allozyme electrophoresis (Polyacrylamide gels and cellulose acetate plates) suggests that queen replacement occasionally occurs. The mean relatedness determined for adult workers in a functionally monogynous population of Leptothorax sp. A was 0.54 and their effective number of mothers therefore 1.5. This is lower than expected and found for monogynous colonies. Dominance rank is apparently not correlated with queen morph, weight, and size, but an influence of insemination, age, or previous reproductive experience is likely.     

Mating flight, metrosis, and semi-claustrality in the seed-harvester ant Pogonomyrmex salinus (Hymenoptera, Formicidae)

Here we examine dispersal, metrosis, and claustrality in the seed-harvester ant Pogonomyrmex salinus at an unusually large mating aggregation. We found that mode of queen dispersal from the mating aggregation is not a function of queen mass and that wing damage among queens was relatively rare. P. salinus is haplometrotic in the field and foundress queens placed together in forced associations eventually fight to the death. While queens of Pogonomyrmex salinus can survive claustrally, producing a single minim from their body reserves in the laboratory, fed queens produce up to four significantly larger minims along with concurrent larvae and pupae during the same period. Since queens forage in the field, we interpret claustrality as a secondary reserve strategy when foraging fails, and suggest that foraging is obligate for P. salinus queens in an overdispersed and temperate environment. Thus, nest founding strategies employed by P. salinus may be environmentally determined and represent a continuum between fully claustral and obligate foraging. We discuss our results with reference to theories of pleometrosis and claustral colony founding. Received 12 November 2004; revised 12 April 2004; accepted 29 July 2005.     

Genetic and social structure of the queen size dimorphic ant Leptothorax cf. andrei

1. The study of queen polymorphisms can provide insight into the evolution of alternative life histories in ants. In this paper, results of morphological, social and genetic investigations of the newly discovered queen size dimorphism in Leptothorax cf. andrei are presented. 2. Queens had a bimodal size distribution, and were classified as large (macrogynes) or small (microgynes) queens. Despite their small size, microgynes had a fully developed external flight apparatus and a functional reproductive tract. 3. Queen morphology was not correlated with colony social structure, and the relatedness among nestmate queens was high, indicating secondary polygyny by re‐adoption of related queens (daughters) into existing colonies. 4. The distribution of microsatellite alleles indicated that there is a genetic separation between macro‐ and micro‐gynes but the two morphs belong to the same species. 5. The results support the hypothesis that microgyny in Leptothorax has not evolved as a specialisation to inter‐ or intra‐specific social parasitism but rather is an adaptation to alternative dispersal behaviour.     

Reproductive Competition in Colonies of the Ant Leptothorax gredleri

Colonies of the ant, Leptothorax (s. str.) gredleri may contain several inseminated female reproductives of which typically only one is laying eggs. Observations suggest that “functional monogyny” is caused by aggressive interactions among nestmate queens. Only the most dominant queen reproduces. Subordinate queens either leave the colony to found their own nests solitarily or by budding, or stay in the nest without reproducing, but may eventually replace the dominant queen. The interrelations of life history of L. gredleri, population structure and habitat characteristics are examined.     

Male parentage and queen mating frequency in the bumblebee <Emphasis Type="Italic">Bombus ignitus</Emphasis> (Hymenoptera: bombinae)

Kin selection theory predicts conflict between queens and workers in the social insect colony with respect to male production. This conflict arises from the haplodiploid system of sex determination in Hymenoptera that creates relatedness asymmetries in which workers are more closely related to the sons of other workers than to those of the queen. In annual hymenopteran societies that are headed by a single queen, the mating frequency of the queen is the only factor that affects the colony kin structure. Therefore, we examined the mating structure of queens and the parentage of males in a monogynous bumblebee, Bombus ignitus, using DNA microsatellites. In the seven colonies that were studied, B. ignitus queens mated once, thereby leading to the prediction of conflict between the queen and workers regarding male production. In each of the five queen-right colonies, the majority of the males (95%) were produced by the colony’s queen. In contrast, workers produced approximately 47% of all the males in two queenless colonies. These results suggest that male production in B. ignitus is a conflict between queen and workers.     

Workers of the primitively eusocial wasp <Emphasis Type="Italic">Ropalidia marginata</Emphasis> do not perceive their queen across a wire mesh partition

Queens of primitively eusocial wasps generally have active and behaviourally dominant queens who use physical aggression to suppress worker reproduction. Although a Ropalidia marginata queen is strikingly docile and behaviourally non-dominant, she is completely successful in maintaining reproductive monopoly. R. marginata queens must achieve such reproductive monopoly by some means other than overt physical aggression. Upon loss or removal of the queen, one of the workers (referred to as the potential queen) becomes extremely aggressive and will eventually go on to become the next queen of the colony, if the original queen is not returned. The fact that potential queens are not discernible in the presence of the queen but become obvious within minutes after removal of the queen raises the question of how workers in general and the potential queens in particular, perceive the presence or absence of their queens. Here, we have conducted experiments in which we separate half of the workers from their queen by a wire mesh screen and study their behavioural response to such separation. We demonstrate that the presence of the queen is not perceived across the wire mesh screen, which suggests that if the queen uses a pheromone to signal her presence, then that pheromone is not very volatile.     

Social organization, reproductive behavior and ecology of Leptothorax acervorum (Hymenoptera, Formicidae) from the Sierra de Albarracin in central Spain

Summary: Leptothorax acervorum, an ant species with holarctic range, occurs in an isolated population in the Spanish Sierra de Albarracin. Dissection of dealate females and laboratory observations revealed that in contrast to other European populations, the colonies are monogynous, with one reproductive queen each and a variable number of virgin or mated dealate but not laying females. Most of the latter probably just hibernate in the mother nests, leaving them in the following spring, but a few remain there for longer time, without reproducing. Such colonies then are functionally monogynous. Alate females exhibit a stationary sexual calling, and mating behavior could be studied in the laboratory. Mated females return to the mother nest where they soon shed wings. When developing fertility before or after hibernation they are evicted from the nests; in nature they probably form daughter colonies. Patchy habitat and rough climatic conditions in the Sierra de Albarracin may be responsible for the particular reproductive behavior of L. acervorum in this area. The generally small size difference between queens and workers in the subgenus Leptothorax entails high costs of dispersal and colony foundation by single queens who have to forage for their first brood. Some kind of dependent colony foundation therefore is frequently met with in the subgenus. Notwithstanding the marked biological and a few slight morphological differences between central European L. acervorum and the Spanish population its taxonomic status as yet is unsettled. We refer to this population provisionally as "L. acervorum Albarracin".     

Determining colony densities in wild honeybee populations (<Emphasis Type="Italic">Apis mellifera</Emphasis>) with linked microsatellite DNA markers

Estimating the population size of social bee colonies in the wild is often difficult because nests are highly cryptic. Because of the honeybee (Apis mellifera) mating behaviour, which is characterized by multiple mating of queens at drone congregation areas (DCA), it is possible to use genotypes of drones caught at these areas to infer the number of colonies in a given region. However, DCAs are difficult to locate and we assess the effectiveness of an alternative sampling technique to determine colony density based on inferring male genotypes from queen offspring. We compare these methods in the same population of wild honeybees, Apis mellifera scutellata. A set of linked microsatellite loci is used to decrease the frequency of recombination among marker loci and therefore increase the precision of the estimates. Estimates of population size obtained through sampling of queen offspring is significantly larger than that obtained by sampling drones at DCAs. This difference may be due to the more extensive flying range of queens compared with drones on mating flights. We estimate that the population size sampled through queen offspring is about double that sampled through drones.     

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.     

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.     

Honey Bee Queens Do Not Count Mates to Assess their Mating Success

The mating system of honey bees (genus Apis) is extremely polyandrous, where reproductive females (queens) typically mate with 12 or more males (drones) during their mating flight(s). The evolutionary implications for hyperpolyandry have been subject to considerable debate and empirical testing because of the need to understand the proximate mechanisms that drive such extreme mating behavior despite the potential costs. The ability of queens to gauge and adjust their reproductive success is therefore important for selection to act on queen mating number at both the evolutionary (colony-level) and proximate (individual-level) timescales. We observed the mating flight activities of 80 queens in their respective mating nucleus hives each with a modified entrance that restricts flight attempts. We also attached a small weight (0, 16, or 38 mg) onto each queen’s thorax as a means of imposing additional flight costs. We then compared queens that were restricted from taking multiple mating flights to those that started oviposition after a single flight for their mating numbers as quantified by microsatellite analyses of their respective worker offspring. We found that neither additional weight nor restricted mating attempts had any significant effect on the effective mating frequencies of the experimental queens during their single mating flight. This observation suggests that queens are not adjusting their nuptial flight activity according to their precise mating number during their flight. These findings provide insights into the proximate regulation of honey bee queen mating behavior and the fitness consequences of hyperpolyandry at the colony level.     

Habitat age, breeding system and kinship in the ant Formica fusca

In polygyne ants (multiple queens per colony) factors that affect the distribution and survival of queens may play a key role in shaping the population-wide mating system and colony kin structure. The aim of this paper was to study the breeding system in two populations of different age in the facultatively polygyne ant Formica fusca. Both the observed numbers of queens, and the relatedness patterns among queens, workers and colony fathers were compared in two adjacent populations (ages 17 years and > 100 years) in Southern Finland. The results showed that both the mating system and colony kin structure differed between the study populations. In the old population the relatedness among workers, queens and colony fathers was high. The queens were also related to their mates, resulting in significant inbreeding in workers, but not in queens. Finally, the number of queens per colony fluctuated between years, suggesting queen turnover, and nest-mate queens shared their reproduction unequally (reproductive skew). In the younger population relatedness among queens and workers was lower than in the old population, and the colony fathers were unrelated. Furthermore, inbreeding was absent, and no conclusive evidence was found for reproductive skew among nest-mate queens. Finally, the number of queens per colony appeared more stable between years, although queen turnover occurred also in this population. The observed differences in dispersal and mating behaviour are discussed in the light of a potential connection between population age and habitat saturation.     

Production of winged and wingless males in the ant, <Emphasis Type="Italic">Cardiocondyla minutior</Emphasis>

Summary Wingless (ergatoid) males of the tramp ant Cardiocondyla minutior attack and kill their young ergatoid rivals and thus attempt to monopolize mating with female sexuals reared in the colony. Because of the different strength of local mate competition in colonies with one or several reproductive queens, we expected the production of new ergatoid males to vary with queen number. Sex ratios were mostly female-biased, but in contrast to the sympatric species C. obscurior (Cremer and Heinze, 2002) neither the percentage of ergatoid males nor of female sexuals among the first 20 sexuals produced varied considerably with queen number. As in C. obscurior, experimental colony fragmentation led to the production of winged males, whereas in unfragmented control colonies only ergatoid males eclosed.Received 3 December 2003; revised 20 February 2004; accepted 23 February 2004.     

A shift in colony founding behaviour in the obligate plant-ant Crematogaster (Decacrema) morphospecies 2

Summary. A shift in colony founding behaviour from single queen (haplometrosis) to multiple queens (pleometrosis) was observed locally in the obligate plant-ant Crematogaster (Decacrema) morphospecies 2, which is associated with Macaranga trees in Borneo. In addition, about a quarter of all mature colonies (27 of 95 trees examined) were found to be multiple queen colonies. They arose either directly from pleometrotic founding colonies or secondarily by adoption of additional queens. Using microsatellite markers, we showed that queens in colonies founded through pleometrosis are unrelated and each queen participates in producing worker offspring, albeit with significant skew in a third of the colonies. In mature polygynous colonies, all resident queens contributed to the production of workers and sexual offspring. Relatedness of queens in mature polygynous colonies was not significantly higher than in foundress associations. We hypothesize that increased nest site limitation in this specific interaction trigger the observed shift in colony founding behaviour. Crematogaster msp. 2 inhabits the light demanding pioneer plant species Macaranga pearsonii that is typical for early successional stages of secondary forests. Thus suitable host-plants for colonisation are abundant for only a short time in highly disturbed sites and become increasingly sparse when secondary forest matures.Received 6 September 2004; revised 29 November; accepted 10 December 2004.     

Independent colony founding by ergatoid queens in the ant genus Pogonomyrmex: queen foraging provides an alternative to dependent colony founding

Ant queens exhibit two primary strategies to initiate nests, independent colony founding (ICF) by solitary queens and dependent colony founding (DCF) when the queen starts a nest with a group of workers that disperse on foot from the parent nest. Numerous ant species have wingless (ergatoid) queens, and it is generally assumed that these species exhibit obligate DCF because their lack of wing musculature provides them with few resources to divert towards producing their first brood of workers. Thus, ICF by ergatoid queens is viewed as maladaptive because these queens need to take additional dangerous foraging trips to garner sufficient food to rear their first brood of workers. Contrary to this prediction, I document ICF by ergatoid queens for three species of harvester ants in the genus Pogonomyrmex (subfamily Myrmicinae), P. cunicularius cunicularius, P. cunicularius pencosensis, and P. huachucanus. Queens of P. huachucanus were obligate foragers, i.e., no minim workers could be produced without external food, and one queen of P. cunicularius pencosensis was observed foraging in the field. Abundant and/or predictable food resources likely select for the evolution of semi-claustral nest founding and ICF by these ergatoid queens. Under these conditions, foraging time would be minimized and the number and size of minim workers would be maximized. These benefits should increase founding success, which could compensate for loss of long-range dispersal. Overall, this study demonstrates that care should be taken before concluding that ant colonies employ DCF based solely on queen morphology.     

Queen fecundity and reproductive skew in the termite Nasutitermes corniger

A major goal of studies on social animals is to understand variation in reproduction within and between groups. We used hierarchical regressions to analyze oviposition rates in the neotropical termite Nasutitermes corniger, a species with both monogynous and polygynous colonies. Queen fecundity was a non-linear, saturating function of queen weight. Greater queen weight was associated with larger nest size and with lower numbers of queens per nest, suggesting competition among queens for resources acquired by the colony. The collective egglaying rate of pairs of queens exceeded that of single queens, but further increases in queen number did not raise total fecundity. Skew in oviposition rates, as quantified by Morisita’s index I _δ, averaged 1.2, indicating inequalities in reproductive rates that are only moderately greater than expected for random apportionment. The leveling off of oviposition with increasing queen weight suggests that it is costly for individual females to produce eggs at high rates, which could favor tolerance of reproduction by other females, reducing reproductive skew. We hypothesize that the incentive to tolerate reproduction by other females is especially pronounced for heavier queens, because these queens are close to the limit of their own reproductive capacity. Consistent with this hypothesis, skew in oviposition rates was inversely related to the mean weight of queens within a nest. Received 8 March 2007; revised 17 September 2007; accepted 3 October 2007.     

Absence of nepotism toward imprisoned young queens during swarming in the honey bee

Nepotism is an important potential conflict in animal societies.However, clear evidence of nepotism in the rearing of queensin social insects is limited and controversial. In the honeybee, Apis mellifera, multiple mating by queens leads to thepresence of many patrilines within each colony. When the coloniesreproduce through swarming, workers rear a number of new queens,only a few of which will ultimately head a colony. Workers canpotentially increase their inclusive fitness by nepotisticallyfavoring full-sister over half-sister queens during the queenrearing and elimination process. Most studies have focused oninteractions between workers and immature queens (eggs and larvae)or adult queens who have exited their queen cells. However,adult queens often remain in their queen cells for up to 1 weekafter emerging from their pupa. In this situation, workers preventthe queens from emerging, feed them, and protect them from otheremerged queens. This stage in queen rearing is therefore onein which nepotism could occur. The current study is the firstto investigate the kinship between workers and adult queenswho have not emerged from their queen cells. We observed thefull suite of behaviors expected during this phase of colonyreproduction. Although there was no evidence for nepotism inthe worker–queen interactions, there was a nonrandom distributionacross patrilines of the workers interacting with the queencells. In addition, in one colony we found differential treatmentof fostered (nonkin)-queen cells.     

QUEEN SIGNALING IN SOCIAL WASPS

Social Hymenoptera are characterized by a reproductive division of labor, whereby queens perform most of the reproduction and workers help to raise her offspring. A long‐lasting debate is whether queens maintain this reproductive dominance by manipulating their daughter workers into remaining sterile (queen control), or if instead queens honestly signal their fertility and workers reproduce according to their own evolutionary incentives (queen signaling). Here, we test these competing hypotheses using data from Vespine wasps. We show that in natural colonies of the Saxon wasp, Dolichovespula saxonica, queens emit reliable chemical cues of their true fertility and that these putative queen signals decrease as the colony develops and worker reproduction increases. Moreover, these putative pheromones of D. saxonica show significant conservation with those of Vespula vulgaris and other Vespinae, thereby arguing against fast evolution of signals as a result of a queen–worker arms race ensuing from queen control. Lastly, levels of worker reproduction in these species correspond well with their average colony kin structures, as predicted by the queen signaling hypothesis but not the queen control hypothesis. Altogether, this correlative yet comprehensive analysis provides compelling evidence that honest signaling explains levels of reproductive division of labor in social wasps.     

Sib‐mating in the ant Plagiolepis pygmaea: adaptative inbreeding?

Multiple functional queens in a colony (polygyny) and multiple mating by queens (polyandry) in social insects challenge kin selection, because they dilute inclusive fitness benefits from helping. Colonies of the ant Plagiolepis pygmaea brash contain several hundreds of multiply mated queens. Yet, within‐colony relatedness remains unexpectedly high. This stems from low male dispersal, extensive mating among relatives and adoption of young queens in the natal colony. We investigated whether inbreeding results from workers expelling foreign males, and/or from preferential mating between related partners. Our data show that workers actively repel unrelated males entering their colony, and that queens preferentially mate with related males. These results are consistent with inclusive fitness being a driving force for inbreeding: by preventing outbreeding, workers reduce erosion of relatedness within colonies due to polygyny and polyandry. That virgin queens mate preferentially with related males could result from a long history of inbreeding, which is expected to reduce depression in species with regular sibmating.     

Genetic evidence for landscape effects on dispersal in the army ant Eciton burchellii

Inhibited dispersal, leading to reduced gene flow, threatens populations with inbreeding depression and local extinction. Fragmentation may be especially detrimental to social insects because inhibited gene flow has important consequences for cooperation and competition within and among colonies. Army ants have winged males and permanently wingless queens; these traits imply male‐biased dispersal. However, army ant colonies are obligately nomadic and have the potential to traverse landscapes. Eciton burchellii, the most regularly nomadic army ant, is a forest interior species: colony raiding activities are limited in the absence of forest cover. To examine whether nomadism and landscape (forest clearing and elevation) affect population genetic structure in a montane E. burchellii population, we reconstructed queen and male genotypes from 25 colonies at seven polymorphic microsatellite loci. Pairwise genetic distances among individuals were compared to pairwise geographical and resistance distances using regressions with permutations, partial Mantel tests and random forests analyses. Although there was no significant spatial genetic structure in queens or males in montane forest, dispersal may be male‐biased. We found significant isolation by landscape resistance for queens based on land cover (forest clearing), but not on elevation. Summed colony emigrations over the lifetime of the queen may contribute to gene flow in this species and forest clearing impedes these movements and subsequent gene dispersal. Further forest cover removal may increasingly inhibit Eciton burchellii colony dispersal. We recommend maintaining habitat connectivity in tropical forests to promote population persistence for this keystone species.