Synchronized locomotion can improve spatial accessibility inside ant colonies.

Synchronization is a conspicuous form of collective behaviour that is of crucial importance in numerous biological systems. Ant colonies from the genera Leptothorax and Temnothorax form small colonies, typically made up of only a few hundred workers, and exhibit a form of synchronized behaviour where workers inside colonies' nests become active together in rhythmic cycles that have a period of approximately 20-200 min. However, it is not currently known if these synchronized rhythms of locomotion confer any functional benefit to colonies. By using a combination of multiple image analysis techniques, we show that inactive Leptothorax ants can act as immobile obstacles to moving ants, and that synchronized activity has the potential to reduce the likelihood that individual ants will encounter regions of immobile obstacles that impede access to portions of the nest. We demonstrate qualitatively similar findings using a computational model of confined active particles with oscillating activity.

Insights into the evolution, biogeography and natural history of the acorn ants, genus Temnothorax Mayr (hymenoptera: Formicidae).

BACKGROUND: Temnothorax (Formicidae: Myrmicinae) is a diverse genus of ants found in a broad spectrum of ecosystems across the northern hemisphere. These diminutive ants have long served as models for social insect behavior, leading to discoveries about social learning and inspiring hypotheses about the process of speciation and the evolution of social parasitism. This genus is highly morphologically and behaviorally diverse, and this has caused a great deal of taxonomic confusion in recent years. Past efforts to estimate the phylogeny of this genus have been limited in taxonomic scope, leaving the broader evolutionary patterns in Temnothorax unclear. To establish the monophyly of Temnothorax, resolve the evolutionary relationships, reconstruct the historical biogeography and investigate trends in the evolution of key traits, I generated, assembled, and analyzed two molecular datasets: a traditional multi-locus Sanger sequencing dataset, and an ultra-conserved element (UCE) dataset. Using maximum likelihood, Bayesian, and summary-coalescent based approaches, I analyzed 22 data subsets consisting of 103 ingroup taxa and a maximum of 1.8 million base pairs in 2485 loci. RESULTS: The results of this study suggest an origin of Temnothorax at the Eocene-Oligocene transition, concerted transitions to arboreal nesting habits in several clades during the Oligocene, coinciding with ancient global cooling, and several convergent origins of social parasitism in the Miocene and Pliocene. As with other Holarctic taxa, Temnothorax has a history of migration across Beringia during the Miocene. CONCLUSIONS: Temnothorax is corroborated as a natural group, and the notion that many of the historical subgeneric and species group concepts are artificial is reinforced. The strict form of Emery's Rule, in which a socially parasitic species is sister to its host species, is not well supported in Temnothorax.

Phylogeography of social polymorphism in a boreo-montane ant.

BACKGROUND: The disjunct distribution of several Palearctic species has been widely shaped by the changes in climatic conditions during the Quaternary. The observed genetic differentiation or reproductive isolation between extant populations may be the outcome of their contemporary geographic separation or reproductive incompatibility due to differences in phenotypic traits which have evolved in isolated refugia. In the boreal ant Leptothorax acervorum, colonies from central and peripheral populations differ in social structure: colonies from Central and Northern Europe may contain several equally reproductive queens (facultative polygyny), while in colonies from peripheral populations in Spain only one the most dominant of several queens lays eggs (functional monogyny). By reconstructing the specie's evolutionary and demographic history in Southwestern Europe we examine whether variation in social organization is associated with restricted gene flow between the two social forms. RESULTS: We show that multi-queen colonies from all so far known inner Iberian populations of L. acervorum are functionally monogynous, whereas multi-queen colonies from all Pyrenean populations are polygynous, like those from other previously studied areas in Europe. Our analyses revealed complex spatial-genetic structure, but no association between spatial-genetic structure and social organization in SW-Europe. The population in the western Pyrenees diverged most strongly from other Iberian populations. Moreover, microsatellite data suggest the occurrence of recent bottlenecks in Pyrenean and inner Iberian populations. CONCLUSIONS: Our study shows a lack of reproductive isolation between the two social forms in SW-Europe. This in turn suggests that demographic and spatial patterns in genetic variation as well as the distribution of social phenotypes are better explained by co-variation with climatic, ecological, and historical factors. Moreover, we for the first time show the existence of substantial spatial-genetic structure in L. acervorum, suggesting the existence of multiple refugia in SW-Europe, including two extra-Mediterranean refugia in France. While gene flow among inner Iberian refugia may have been larger during the late glacial, extra-Mediterranean refugia in southern France may have contributed to the post-glacial recolonization of W-Europe.

Noise resistant synchronization and collective rhythm switching in a model of animal group locomotion.

Biology is suffused with rhythmic behaviour, and interacting biological oscillators often synchronize their rhythms with one another. Colonies of some ant species are able to synchronize their activity to fall into coherent bursts, but models of this phenomenon have neglected the potential effects of intrinsic noise and interspecific differences in individual-level behaviour. We investigated the individual and collective activity patterns of two Leptothorax ant species. We show that in one species (Leptothorax sp. W), ants converge onto rhythmic cycles of synchronized collective activity with a period of about 20 min. A second species (Leptothorax crassipilis) exhibits more complex collective dynamics, where dominant collective cycle periods range from 16 min to 2.8 h. Recordings that last 35 h reveal that, in both species, the same colony can exhibit multiple oscillation frequencies. We observe that workers of both species can be stimulated by nest-mates to become active after a refractory resting period, but the durations of refractory periods differ between the species and can be highly variable. We model the emergence of synchronized rhythms using an agent-based model informed by our empirical data. This simple model successfully generates synchronized group oscillations despite the addition of noise to ants' refractory periods. We also find that adding noise reduces the likelihood that the model will spontaneously switch between distinct collective cycle frequencies.

Queen-worker ratio affects reproductive skew in a socially polymorphic ant.

The partitioning of reproduction among individuals in communally breeding animals varies greatly among species, from the monopolization of reproduction (high reproductive skew) to similar contribution to the offspring in others (low skew). Reproductive skew models explain how relatedness or ecological constraints affect the magnitude of reproductive skew. They typically assume that individuals are capable of flexibly reacting to social and environmental changes. Most models predict a decrease of skew when benefits of staying in the group are reduced. In the ant Leptothorax acervorum, queens in colonies from marginal habitats form dominance hierarchies and only the top-ranking queen lays eggs ("functional monogyny"). In contrast, queens in colonies from extended coniferous forests throughout the Palaearctic rarely interact aggressively and all lay eggs ("polygyny"). An experimental increase of queen:worker ratios in colonies from low-skew populations elicits queen-queen aggression similar to that in functionally monogynous populations. Here, we show that this manipulation also results in increased reproductive inequalities among queens. Queens from natural overwintering colonies differed in the number of developing oocytes in their ovaries. These differences were greatly augmented in queens from colonies with increased queen:worker ratios relative to colonies with a low queen:worker ratio. As assumed by models of reproductive skew, L. acervorum colonies thus appear to be capable of flexibly adjusting reproductive skew to social conditions, yet in the opposite way than predicted by most models.

Avoid mistakes when choosing a new home: Nest choice and adoption of Leptothorax ant queens.

In ants, mating and colony founding are critical steps in the life of ant queens. Outside of their nests, young queens are exposed to intense predation. Therefore, they are expected to have evolved behavior to accurately and quickly locate a nesting place. However, data on the early life history of female reproductives are still lacking. Leptothorax gredleri is a suitable model organism to study the behavior of young queens. Reproductives can be reared under artificial conditions and readily mate in the laboratory. After mating, L. gredleri queens have the options to found solitarily, seek adoption into another colony, or return into their natal nest. In this study, we investigated the decision-making processes of female sexuals before and after mating. In particular, we tested whether female sexuals use chemical cues to find their way back to the nest, studied if they prefer their own nest over other nesting sites and followed the adoption dynamics of mated queens over 8 weeks (plus hibernation and spring). We showed that female sexuals and freshly mated queens spent more time on substrate previously used by workers from their own colony and from another colony than on a blank substrate. This discriminatory capability of queens appears to be lost in old, reproductive queens. Nest choice experiments showed that female sexuals and freshly mated queens can distinguish their own nest while old mated queens do not. When reintroduced in their maternal colony, young queens were readily adopted, but a few weeks later aggression against young queens led to their emigration from the maternal nest and eventually also death.

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.