Alternative adaptations, sympatric speciation and the evolution of parasitic, inquiline ants

Inquiline ant species are workerless social parasites whose queens reproduce in colonies of other species alongside the host queens. Inquilines arise either when one non-parasitic species evolves into an inquiline parasite of another non-parasitic species (the interspecific hypothesis), or by the speciation of intraspecific inquilines from their host stock (the intraspecific hypothesis): it is unlikely that inquilines evolve from other forms of social parasite. This paper reviews the evidence for and against the inter-and intraspecific hypotheses. All inquilines are close phylogenetic relatives of their host species (loose ‘Emery's rule’), and some are their host's closest relative (strict ‘Emery's rule’). A problem for the interspecific hypothesis is how to explain the strict Emery's rule, because phylogenetic constraints on host choice are probably quite weak. By contrast, the intraspecific hypothesis has difficulty accounting for the parasites' sympatric reproductive isolation. Facultative polygyny, in which queens may found colonies alone or by adoption into an existing multi-queen colony, should promote the evolution of small intraspecific inquilines. This is because small colony-founding queens should preferentially seek adoption, which provides the opportunity to produce a sexual-only brood. We suggest that microgynes, i.e. miniature queens found in some polygynous ants, represent such parasites. We review the evidence that inquiline species have evolved intraspecifically from microgynes in Myrmica ants. The coexistence within a species of a monogynous (singly-queened) and a polygynous form is probably a phenomenon usually unconnected with inquiline evolution. The reproductive isolation of intraspecific inquilines plausibly arises from divergent breeding behaviour associated with the parasites' small size. Such divergence could involve either a temporal separation in mating episodes, with small parasites maturing early, or a spatial separation, with small males being sexually-selected to mate near the nest with small queens seeking adoption, instead of in mating aggregations. We conclude that inquiline species strictly following Emery's rule could have evolved by the intraspecific route. If so, such species provide evidence for West-Eberhard's “alternative adaptation” hypothesis that between-species diversity frequently stems from diversity within species. They also represent likely cases of sympatric speciation. We suggest work on the parasites' phytogeny, genetics, behaviour and mating biology to test these conclusions further.     

Habitat differences and occurrence of native and exotic ants on Okinawa Island

Ant fauna on Mallotus japonicus, a shrub with extrafloral nectaries, was investigated in two types of habitat (the villages and the forest‐edges) on subtropical Okinawa Island, Japan. Twenty and 16 ant species were found, including 11 and 6 tramp species in the villages and in the forest‐edges, respectively. Occurrence of tramp species was higher in the villages than in the forest edges, supporting the idea that tramp species tend to dominate in disturbed habitats. Nevertheless, the richness of native ants was almost the same across the two habitat types. Consequently, the ant species diversity on M. japonicus appeared higher in the villages. However, monitoring has to be continued to determine whether the above findings represent a stable phenomenon of the community or just a temporary state.     

Evolutionary transition to a semelparous life history in the socially parasitic ant Acromyrmex insinuator

The recently discovered social parasite Acromyrmex insinuator ( 25 ) exploits colonies of the leafcutter ant A. echinatior. We document that A. insinuator represents a rare early stage in the evolution of social parasitism, because a worker caste is still partially present and mating phenology has remained at least partially similar to that of the host. A. insinuator is tolerant of host queens, and sexual offspring produced in parasitized colonies can be either exclusively A. insinuator or a mix of A. insinuator and A. echinatior. The remarkably high abundance of A. insinuator in nests of the investigated Panamanian host population and the fact that A. insinuator colonies readily reproduce under laboratory conditions allowed us to test evolutionary predictions on reproductive life history evolution that are not possible in most other socially parasitic ants. We show that (1) A. insinuator has a semelparous ‘big bang’ reproductive life history which exploits host colonies without leaving reserves for survival; (2) social parasite sexuals are significantly smaller than A. echinatior host sexuals, but still large compared to host workers, confirming an evolutionary scenario of gradual size reduction and loss of the worker caste after transition towards a socially parasitic life history; (3) major changes in the life history of ants can evolve relatively quickly compared to adaptations in morphology, caste differentiation and mating phenology.     

Testes degeneration in ants: a histological study of Gnamptogenys bicolor

Allard, D., Ito, F., Aikawa, Y., Gotoh, A. and Billen, J. 2010. Testes degeneration in ants: a histological study of Gnamptogenys bicolor. —Acta Zoologica (Stockholm) 92 : 372–376. Sperm synthesis in the males of social Hymenoptera is not continuous, as the testes degenerate before the males become sexually active. This article presents the first histological study focusing on the degeneration of testes tissue, using adult males of the ant Gnamptogenys bicolor. At eclosion, spermatogenesis is still in process in the apical part of the testes, while the basal spermatocysts are all packed with mature sperm. During the second week after eclosion, the first signs of degeneration appear as small clear vacuoles inside the spermatocysts, interspersed between maturing sperm cells. Sperm starts migrating into the vasa deferentia shortly after. As spermatocysts void their contents into the vasa deferentia, the follicles start shrinking and losing their shape and the fibrillar capsule tissue gets looser. Cellular breakdown yields dark vacuoles of various density and large granular, multi‐layered bodies.     

Abundance and distribution of organic mound-building ants of the Formica rufa group in Yellowstone National Park

Red wood ants (Formica rufa group) are ubiquitous in many conifer and mixed‐conifer forests of northern Europe and Asia. In contrast, relatively little is known about the abundance and distribution of the 24 North American F. rufa group species. As ants are important components of most soil invertebrate communities and are considered ecosystem engineers that alter the flow of energy and nutrients through terrestrial systems, it is important to gain information on their distribution and abundance. We conducted a survey for red wood ant mounds in Yellowstone National Park (YNP), Wyoming/Montana, USA, where human disturbance has been kept to a minimum for over 130 years. We found a total of 85 red wood ant mounds (0.11 to 0.17 mounds/ha) on 327 km of roads and 180 km of the hiking trails we surveyed. The occurrence of ant mounds was higher then expected by random distribution at elevations between 1600 and 2400 m, annual precipitation of 250 to 760 mm, middle and late successional lodgepole pine, late successional Douglas fir forest and non‐forested grassland/sagebrush prairie vegetation. Additionally, mounds were clustered in gently sloped not north‐exposed locations and in areas that had not recently burned. Most of the mounds detected were inhabited by Formica obscuripes Forel, which occupied 94% of the mounds sampled. Based on a multi‐criteria binary Geographic Information System model that we developed, we found that ant mounds were to be expected with a high probability in less then 1% of the YNP area. These results together with the detected low density and small size of the red wood ant mounds within the study area suggest that these insects have a much lower impact on invertebrate biodiversity and ecosystem processes, such as forest productivity and carbon and nutrient cycling on the ecosystem scale compared with their counterparts in European or Asian systems.     

Rareness of certain Mediterranean ant species: fact or artifact?

Summary A checklist of the ants recorded during five years in a small urban area (400 m²) in Sant Cugat (Barcelona, Spain) is presented. 23 species nest in the area, 7 species (hypogaeic or parasitic) probably nest and 5 species nest in the vicinity; five additional species have also been recovered from the area. The rarity of some of these (Leptanilla sp.,Hypoponera abeillei (Emery),Cryptopone ochraceum (Mayr),Smithistruma tenuipilis (Emery),Epitritus argiolus Emery,Trichoscapa membranifera Emery,Chalepoxenus muellerianus (Finzi),Anergates atratulus (Schenck),Plagiolepis xene (Schenck),Lasius carniolicus Mayr) and the high number of species suggest that a) rarity in Mediterranean ant species may be explained by the insufficient sampling in time of adequate microhabitats (hypogaeic species) or by the inconspicuousness of social parasitic species, and b) that previously published species lists of ants from Mediterranean habitats might be rather incomplete.     

Comparison of the ant communities of annually inundated and terra firme forests at Trombetas in the Brazilian Amazon

Summary The composition of the ant community was assessed along standardized 100 m transects in annually flooded Varzea forest and in terra firme forests on sandy soil (Flanco forest) and on claytopped mesas (Planalto forest). Standardized samples were taken by unit-time hand collecting (day and night times), sweeping, beating, baiting and by Winkler sacks. A total of 156 species, representing 49 genera were found, of which 98, 88 and 55 were respectively found in the Planalto, Flanco and Varzea forests. Species lists are presented and the ant community composition and species richness are compared between the three forests. By considering the nesting and foraging habits of the various species, the differences in overall community composition are related to the forest type and susceptibility to inundation of the three forests which were surveyed.The data confirm the view that tropical rain forests support an extremely diverse ant fauna and comparisons with other forested areas suggest that ant species richness declines in subtropical and temperate rain forests. Although alpha diversity is high, species turnover between forests is lower than expected, suggesting that ant species richness in this forested region is not as great as is implied in some published estimates of global arthropod diversity.     

Macroevolutionary patterns in the origin of mutualisms involving ants

Ants are a diverse and abundant insect group that form mutualistic associations with a number of different organisms from fungi to insects and plants. Here, we use a phylogenetic approach to identify ecological factors that explain macroevolutionary trends in the mutualism between ants and honeydew-producing Homoptera. We also consider association between ant-Homoptera, ant-fungi and ant-plant mutualisms. Homoptera-tending ants are more likely to be forest dwelling, polygynous, ecologically dominant and arboreal nesting with large colonies of 10(4)-10(5) individuals. Mutualistic ants (including those that garden fungi and inhabit ant-plants) are found in under half of the formicid subfamilies. At the genus level, however, we find a negative association between ant-Homoptera and ant-fungi mutualisms, whereas there is a positive association between ant-Homoptera and ant-plant mutualisms. We suggest that species can only specialize in multiple mutualisms simultaneously when there is no trade-off in requirements from the different partners and no redundancy of rewards.