Queen Morphs Research Articles

Social organization and the evolution of life-history traits in two queen morphs of the ant Temnothorax rugatulus.

During the evolution of social insects, not only did life-history traits diverge, with queens becoming highly fecund and long lived compared with their sterile workers, but also individual traits lost their importance compared with colony-level traits. In solitary animals, fecundity is largely influenced by female size, whereas in eusocial insects, colony size and queen number can affect the egg-laying rate. Here, we focused on the ant Temnothorax rugatulus, which exhibits two queen morphs varying in size and reproductive strategy, correlating with their colony's social organization. We experimentally tested the influence of social structure, colony and body size on queen fecundity and investigated links between body size, metabolic rate and survival under paraquat-induced oxidative stress. To gain insight into the molecular physiology underlying the alternative reproductive strategies, we analysed fat body transcriptomes. Per-queen egg production was lower in polygynous colonies when fecundity was limited by worker care. Colony size was a determinant of fecundity rather than body size or queen number, highlighting the super-organismal properties of these societies. The smaller microgynes were more frequently fed by workers and exhibited an increase in metabolic activity, yet they were similarly resistant to oxidative stress. Small queens differentially expressed metabolic genes in the fat body, indicating that shifts in molecular physiology and resource availability allow microgyne queens to compensate for their small size with a more active metabolism without paying increased mortality costs. We provide novel insights into how life-history traits and their associations were modified during social evolution and adapted to queen reproductive strategies.

Life‐history trait variation in a queen‐size dimorphic ant

1. Size polymorphism is often connected to alternative life‐history traits, which may eventually lead to distinct size classes. In the ant Myrmica ruginodis, larger macrogyne and smaller microgyne queen morphs have been suggested to follow different reproductive strategies, which has presumably resulted in several differences in their key life‐history traits.2. In this study, we examine the association of queen‐size morphs with colony queen number (monogyny vs. polygyny), dispersal and queen recruitment patterns, as well as habitat associations of the queen morphs. We do this by sampling established queens from a large number of excavated nests from several populations, estimating genetic relatedness among coexisting queens and pitfall trapping free‐ranging wingless queens.3. Our results show that associations of queen morphs with colony queen number and nest‐founding strategy holds only partly. The morph frequencies vary widely across populations from practically pure macrogyne to more than 50% microgyne, but the expected association of macrogyne occurrence with monogyny and microgyne with polygyny is not universal. Dispersal and queen recruitment patterns also show that although most macrogynes participate in nuptial flights and most microgynes are recruited back to their natal nests, a fraction of both morphs use the alternative strategy.4. The polygynous microgyne morph has been suggested to specialize in stable habitats, but our results from Finnish mesic heath forests do not support this. This study shows that factors other than just queen size also influence life‐history trait variation and reproductive strategies in ants.

The dimorphic queens of Polyrhachis australis (Hymenoptera: Formicidae)

AbstractThe formicine weaver ant Polyrhachis australis Mayr is one among many species of ants that have queens of two more or less distinct sizes, usually referred to as microgynes and macrogynes. The current consensus explains this phenomenon in terms of alternative strategies of dispersal and reproduction: microgynes typically walk to found new nests locally, macrogynes typically fly to found new nests farther afield. This study addressed four questions bearing on that interpretation: is the dimorphism complete (without intermediate sizes) or continuous in P. australis? Does the proportion of macrogynes correlate with the number of queens in the nest? Does the microgyne/macrogyne proportion vary over time? And are brachypterous individuals always microgynes? The implications of these questions concern the fitness status of the intermediate size individuals, the relative fitnesses of the two main morphs as the number of queens diminishes, the influence of habitat conditions on the production of the morphs, and the dispersal behaviour of microgynes. The size distribution of P. australis queens was incompletely bimodal, with some individuals of intermediate size. The proportion of macrogynes in nests increased with decreasing numbers of queens in nests, suggesting that macrogynes do better in whatever contests take place between queens during the ergonomic stage. Between 2009 and 2014, the proportion of macrogynes in nests fell from about 1 in 3 or 4, to less than 1 in 11, perhaps reflecting a differential production of queen morphs in saturated and unsaturated habitats: it profits a colony to produce more microgynes and fewer macrogynes when local conditions favour the easy founding of new nests by walking. Brachyptery was centred on the smallest microgynes, consistent with their assumed mode of dispersal for which wings are not needed.

Genetic divergence between the sympatric queen morphs of the ant Myrmica rubra.

Pairs of obligate social parasites and their hosts, where some of the parasites have recently diverged from their host through intraspecific social parasitism, provide intriguing systems for studying the modes and processes of speciation. Such speciation, probably in sympatry, has also been propounded in the ant Myrmica rubra and its intraspecific social parasite. In this species, parasitism is associated with queen size dimorphism, and the small microgyne has become a social parasite of the large macrogyne. Here, we investigated the genetic divergence of the host and the parasite queen morphs in 11 localities in southern Finland, using nuclear and mitochondrial markers of queens and workers. We formulated and tested four speciation-related hypotheses that differed in the degree of genetic divergence between the morphs. The queen morphs were genetically distinct from each other with little hybridization. In the nuclear data, when localities were nested within queen morphs in the hierarchical amova, 39% of the genetic variation was explained by the queen morph (standardized F'CT = 0.63, uncorrected FCT = 0.39), whereas 18% was explained by the locality (F'SC = 0.39, FSC = 0.29). This result corroborated the hypothesis of advanced sympatric speciation. In contrast, the mitochondrial DNA could not settle between the hierarchical levels of locality and queen morph, thus substantiating equally the hypotheses of incipient and advanced sympatric speciation. Together, our results support the view that the microgynous parasite has genetically diverged from its macrogynous host to the level of a nascent species.

Phylogeography of the antMyrmica rubraand its inquiline social parasite

Widely distributed Palearctic insects are ideal to study phylogeographic patterns owing to their high potential to survive in many Pleistocene refugia and—after the glaciation—to recolonize vast, continuous areas. Nevertheless, such species have received little phylogeographic attention. Here, we investigated the Pleistocene refugia and subsequent postglacial colonization of the common, abundant, and widely distributed ant Myrmica rubra over most of its Palearctic area, using mitochondrial DNA (mtDNA). The western and eastern populations of M. rubra belonged predominantly to separate haplogroups, which formed a broad secondary contact zone in Central Europe. The distribution of genetic diversity and haplogroups implied that M. rubra survived the last glaciation in multiple refugia located over an extensive area from Iberia in the west to Siberia in the east, and colonized its present areas of distribution along several routes. The matrilineal genetic structure of M. rubra was probably formed during the last glaciation and subsequent postglacial expansion. Additionally, because M. rubra has two queen morphs, the obligately socially parasitic microgyne and its macrogyne host, we tested the suggested speciation of the parasite. Locally, the parasite and host usually belonged to the same haplogroup but differed in haplotype frequencies. This indicates that genetic differentiation between the morphs is a universal pattern and thus incipient, sympatric speciation of the parasite from its host is possible. If speciation is taking place, however, it is not yet visible as lineage sorting of the mtDNA between the morphs.

Low queen mating frequency in the seed-harvester ant Pogonomyrmex (Ephebomyrmex) pima: implications for the evolution of polyandry

The evolution of polyandry is a central problem in the study of insect mating systems, and both material and genetic benefits have been proposed to offset the presumed costs of multiple mating. Although most eusocial Hymenoptera queens mate with just one or occasionally two males, high levels of polyandry are exhibited by several taxa, including seed-harvester ants of the genus Pogonomyrmex. Previous studies of queen mating frequency in Pogonomyrmex have focused on monogynous (one queen per colony) species in the subgenus Pogonomyrmex. We performed a genetic mother–offspring analysis of mating frequency in Pogonomyrmex (Ephebomyrmex) pima, a queen-dimorphic species with dealate and intermorph queens that differ in colony structure (intermorph colonies contain multiple queens). Our results demonstrate that both dealate and intermorph queens of P. (E.) pima are typically single maters, unlike their congeners analyzed thus far. Polyandry appears to be a derived trait in Pogonomyrmex, but comparative tests between P. (E.) pima queen morphs and across the genus provide no evidence that it evolved as an adaptation to increase genetic diversity within colonies or to obtain more sperm, respectively.

Social and genetic structure of a supercolonial weaver ant, Polyrhachis robsoni, with dimorphic queens

We studied a population of the Australian weaver ant Polyrachis robsoni with regard to variation in the morphology of its winged queens using six newlydeveloped microsatellite markers. Morphometrically the queens fell clearly into two groups, macrogynes and microgynes, with the latter an isometric reduction of the former. Aggression tests showed that hostility between ants from different nests was minimal. Nests frequently contained numbers of both queen types, with microgynes about twice as numerous as macrogynes. Nestmate workers, microgynes, and macrogynes, were significantly related to others within their caste, with macrogynes more highly related than the other castes. Relatedness values between these groups of nestmates were also significant. Pairwise relatedness values were consistent with both queen morphs producing workers. At the population level, microgynes from different nests were also significantly related and there was a weak inverse relationship between pairwise relatedness value between individuals and distance between nests.We conclude that this species is supercolonial and that the two queen morphs are part of the same population.

Three queen morphs with alternative nest-founding behaviors in the ant, Temnothorax longispinosus

In many ant species, multiple modes of founding new colonies occur in the same population. These modes include dependent founding, independent founding by haplometrosis (single queen), and independent founding by pleometrosis (multiple queens). In several cases, a dimorphism in queen size has been found, such that each morph specializes in a particular nest-founding behavior. I investigated queen size in the ant Temnothorax longispinosus in several southern Wisconsin populations and found three distinct queen morphs: small queens with very low fat content and short wings, large queens with low fat and long wings, and large queens with high fat and very long wings. Several traits associated with founding behavior correlated with these queen sizes. Small queens were produced in lower numbers, were more common in polygynous nests, and returned to the nest in higher proportions than both large queen morphs. The size ranges and fat levels of each queen morph were similar to those of other species that specialize in either haplometrosis (very large, high fat), pleometrosis (large, low fat), or dependent founding (small and low fat). However, there was extensive overlap in several of the founding behaviors, suggesting that the morphs in these populations have some flexibility in founding behavior. The queen morphs in these populations of T. longispinosus may resemble early stages in the evolution of more specialized dispersal polymorphisms found in other ant species.

Queen polymorphism and queen-morph related facultative polygyny in the ant, Myrmecina graminicola (Hymenoptera, Formicidae)

Polymorphism of the functional queens in Myrmecina graminicola is analyzed. Both gynomorphs (G-♀♀; G) and a wide range of intermorphs (I-♀♀; I) occur, which all are usually mated and egg-laying. Colonies having a gynomorphic queen are always monogynous, whereas about 57% of all colonies with intermorphic queens are polygynous, having two or more coexisting functional queens. The female sexual offspring of individual gynomorphic queens either consists of gynomorphs only, or exclusively of intermorphs. Intermorphic queens may have exclusively intermorphic female sexual progeny, or simultaneously both gynomorphs and intermorphs. Single colonies in laboratory culture produce the same kind of female progeny over several subsequent breeding cycles (artificially compressed "years" of 9–10 months). No environmental influence on queen morph determination could be detected. A genetically mediated queen polymorphism, as in Harpagoxenus sublaevis and Leptothorax sp. A, is suggested. Colony sizes vary considerably, with polygynous I-queen colonies being largest (57.2 ± 34.3 s.d. workers), followed by G-queen colonies (44.6 ± 22.7 s.d.) and monogynous I-queen colonies (34.4 ± 23.7 s.d.), suggesting occasional budding of polygynous colonies.

Queen polymorphism in the Australian ant Monomorium sp.10

The Australian ant Monomorium sp.10 exhibits a queen polymorphism with initially winged gynomorphs and wingless intermorphs. Female sexuals eclosing from brood produced by a gynomorphic queen are always winged, whereas offspring of intermorphic queens consists of both winged and wingless female sexuals. The exchange of brood between colonies suggests that queen morph is determined either genetically or by a maternal effect. We discuss the occurrence of genetically based queen polymorphism in ants and its ecological significance.

Size-dimorphism in the queens of the North American ant Leptothorax rugatulus (Emery)

With recent findings of an unexpected variability in the reproductive behaviour of ant sexuals, their morphology has become an area of major evolutionary interest (Heinze and Tsuji, 1995). Here we report on the occurrence of two queen morphs in Leptothorax rugatulus (Hym., Formicidae): Microgynes (small queens), exceeding worker-size only marginally, and macrogynes, which are, typically for the subgenus Myrafant, about twice as big as their workers. The frequency distribution of queen-size is clearly bimodal, in contrast to worker- and male-size. The average size of queens is highly correlated with the size of daughters in field-collected colonies, whereas within colonies no correlation between the average queen-size and the size of workers or males exists. This gives additional support that size-dimorphism is due to a specific, transmissible size reduction of the microgynes which could be based on genetics, the environment or both. This reduction is quasi-isometric, with a slightly smaller thorax-to-head ratio in microgynes, and scanning electron microscopy does not reveal any significant degeneration of the pterothorax, ocelli or number of ommatidia. The frequency of microgynes at different sample sites is highly variable, correlating well with the prevailing social structure in the respective subpopulations. Indeed, the majority of macrogynes is found in monogynous colonies, while microgynes abound in polygynous ones, which is strong evidence for an alternative dispersal tactic. However, the expected correlation to altitude or latitude was not found and further investigations are needed to reveal proximate and ultimate causes of this prevalent polymorphism between two types of female ant reproductives.

Dynamics and Consequences of Hierarchy Formation in the AntLeptothoraxsp. A

AbstractIn the Nearctic antLeptothoraxsp. 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 ofLeptothoraxsp. 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.

Queen Size Variation in the Ponerine AntPonera coarctata(Hymenoptera: Formicidae)

Queens ofPonera coarctatashow a pronounced variation in size as measured by ommatidia number and Weber's alitrunk length. Isometric size variation and the normal distribution of size categories indicate that, despite these differences, only one queen morph exists. Queen size varies less within colonies than between colonies, and thus appears to be colony specific. Ovary length apparently varies with queen size. Similar size variations as in queens also occured in males, but not in workers.