Biogeography of mutualistic fungi cultivated by leafcutter ants.

Leafcutter ants propagate co-evolving fungi for food. The nearly 50 species of leafcutter ants (Atta, Acromyrmex) range from Argentina to the United States, with the greatest species diversity in southern South America. We elucidate the biogeography of fungi cultivated by leafcutter ants using DNA sequence and microsatellite-marker analyses of 474 cultivars collected across the leafcutter range. Fungal cultivars belong to two clades (Clade-A and Clade-B). The dominant and widespread Clade-A cultivars form three genotype clusters, with their relative prevalence corresponding to southern South America, northern South America, Central and North America. Admixture between Clade-A populations supports genetic exchange within a single species, Leucocoprinus gongylophorus. Some leafcutter species that cut grass as fungicultural substrate are specialized to cultivate Clade-B fungi, whereas leafcutters preferring dicot plants appear specialized on Clade-A fungi. Cultivar sharing between sympatric leafcutter species occurs frequently such that cultivars of Atta are not distinct from those of Acromyrmex. Leafcutters specialized on Clade-B fungi occur only in South America. Diversity of Clade-A fungi is greatest in South America, but minimal in Central and North America. Maximum cultivar diversity in South America is predicted by the Kusnezov-Fowler hypothesis that leafcutter ants originated in subtropical South America and only dicot-specialized leafcutter ants migrated out of South America, but the cultivar diversity becomes also compatible with a recently proposed hypothesis of a Central American origin by postulating that leafcutter ants acquired novel cultivars many times from other nonleafcutter fungus-growing ants during their migrations from Central America across South America. We evaluate these biogeographic hypotheses in the light of estimated dates for the origins of leafcutter ants and their cultivars.

A new ant genus from southern Argentina and southern Chile, Patagonomyrmex (Hymenoptera: Formicidae).

The ant genus Pogonomyrmex (Hymenoptera: Formicidae: Myrmicinae) comprises 71 described species that occur in North America, South America, and Hispaniola, and it is the nominal genus in the recently established tribe Pogonomyrmecini. A molecular phylogeny using 3,647 base pairs from fragments of one mitochondrial gene (cytochrome oxidase I) and five nuclear genes (long-wavelength rhodopsin, elongation factor 1α F1, elongation factor 1α F2, wingless, rudimentary) inferred that Pogonomymrex was not monophyletic. The vast majority of species belonged to a monophyletic clade (Pogonomyrmex sensu stricto), whereas species in the Po. angustus-group formed a second lineage outside of Pogonomyrmex and Hylomyrma, the latter being the only other genus in the tribe. To maintain monophyly of Pogonomyrmex, we create the genus Patagonomyrmex gen. n., which consists of the three angustus-group species (Patagonomyrmex angustus comb. n., Patagonomyrmex laevigatus comb. n., and Patagonomyrmex odoratus comb. n.) that are sister to all other pogonomyrmecines. The genus is restricted to southern Argentina and southern and southcentral Chile. Workers are characterized by: (1) poorly-developed psammophore that consists of short to medium-length hairs scattered over the ventral surface of the head, (2) anterior clypeal margin strongly convex in full-face view, (3) clypeus strongly convex in profile, (4) superior and inferior propodeal spines long, (5) dorsal surface of promesonotum smooth and shiny, lacking rugae or if rugae present, then interrugae strongly granulate-punctate, (6) anteroventral margin of peduncle of the petiole with a small, acuminate spine, and (7) palp formula 5,4. We also provide information on biology, distribution maps, and a key to workers, queens, and males in English and Spanish.

A taxonomic revision of South American species of the seed-harvester ant genus Pogonomyrmex (Hymenoptera: Formicidae). Part I.

South American species in the seed-harvester ant genus Pogonomyrmex (subfamily Myrmicinae) are interesting biologically because of their numerous queen phenotypes and life histories. This paper provides a taxonomic revision and reviews the natural history for 21 South American species of Pogonomyrmex so that we can better study their rich and interesting ecology, life history, and evolution. Species treated herein comprise all South American species-groups except for the brevibarbis and rastratus-groups. The following taxa are raised to species: pencosensis Forel 1914 and serpens Santschi 1922. The following new synonomies are proposed: bruchi Forel 1913 is synonomized under coarctatus Mayr 1868 and cunicularius carnivora Santschi 1925 under serpens Santschi 1922. The following new species is described: tinogasta. This paper redescribes workers of all species, and I describe queens and diagnose males for the following species: bispinosus (ergatoid queen, male), inermis (queen, male), laticeps (male), lobatus (queen, male), micans (queen), naegelii (ergatoid queen), pencosensis (ergatoid queen), serpens (ergatoid queen), tinogasta (brachypterous queen), and uruguayensis (queen, male). A neotype was designated for the untraceable or possibly lost type of P. bispinosus, and a holotype or lectotype was designated from syntypes for all other previously described taxa in order to provide a single name-bearing specimen and to facilitate future taxonomic studies. Of the 21 species treated herein, five species have ergatoid (wingless) queens (bispinosus, cunicularius, pencosensis, serpens, mayri), two have brachypterous (short-winged) queens (mendozanus, tinogasta), and two have dimorphic queens (winged and ergatoid in naegelii, brachypterous and ergatoid in laticeps). I also provide keys for workers and queens (in English and Spanish), photographs of all castes, distribution maps, and a summary of known biology.

A New Species of Seed-harvester Ant, Pogonomyrmex hoelldobleri (Hymenoptera: Formicidae), from the Mohave and Sonoran Deserts of North America.

Pogonomyrmex magnacanthus Cole was described as a distinct species; unusually large eyes and a high ocular index (maximum eye diameter/head width) were listed as diagnostic characters. However, examination of numerous series of P. magnacanthus revealed that both characters were highly variable, and that these series consisted of P. magnacanthus plus an undescribed species, Pogonomyrmex hoelldobleri Johnson, Overson & Moreau sp. nov. This paper describes all three castes of P. hoelldobleri as well as the alate queen of P mohavensis, which is very similar to that of P. hoelldobleri. A molecular phylogeny that consisted of 3,703 bp from one mitochondrial and five nuclear gene fragments-supported the monophyly of P. hoelldobleri, P. magnacanthus, and P mohavensis. Pogonomyrnex inagnacanthus can be separated from other P. californicus group species based on: (1) its unusually large eyes, (2) a high ocular index, and (3) a malar ratio that is typically < or = 1.0. Pogonomyrmex hoelldobleri can be separated from other P. californicus group species based on the combination of: (1) eyes not unusually large, (2) cephalic rugae not forming circumocular whorls, but rather converging posterior to the eyes, usually near the vertex, (3) mandible with seven teeth, and (4) interrugal spaces on pronotal sides moderately to strongly granulate, dull to weakly shining. Pogonomyrmex mohavensis can be separated from other P. californicus group species based on the combination of: (1) eyes not unusually large, (2) cephalic rugae not forming circumocular whorls, but rather extending more or less directly to the vertex or converging only slightly near the vertex, (3) mandible with six teeth (a seventh sometimes occurs as a denticle between the basal and sub-basal teeth), and (4) interrugal spaces on pronotal sides smooth and shining to weakly punctate and moderately shining. We also provide field observations and distribution maps for P. magnacanthus, P. hoelldobleri, and P. mohavensis, and an updated key to P californicus group species that occur in central and western North America.

Selection against aerial dispersal in ants: two non-flying queen phenotypes in Pogonomyrmex laticeps.

The South American seed-harvester ant Pogonomyrmex laticeps has dimorphic queens: ergatoid (permanently wingless) and brachypterous (short, non-functional wings). Surveys in western Argentina indicated that colonies near Chilecito, La Rioja Province, produced only ergatoid queens, while those near Punta Balasto, Catamarca Province (263 km away), produced only brachypterous queens. Brachypterous queens were significantly larger than ergatoid queens for 10 of 11 external characters, but both phenotypes had comparable reproductive potential, i.e., a spermatheca and a similar number of ovarioles. Using normal winged queens of the closely related P. uruguayensis for comparison, we determined that both queen phenotypes in P. laticeps had a full set of dorsal thoracic sclerites, albeit each sclerite was much reduced, whereas workers had a thorax without distinct dorsal sclerites. Sclerites were fused and immobile in ergatoid queens, while they were separable and fully articulated in brachypterous queens. Both phenotypes lacked the big indirect flight muscles, but brachypterous queens retained the tiny direct flight muscles. Overall, this dimorphism across populations indicates that there are alternative solutions to selective pressures against flying queens. We lack field data about colony founding strategy (independent or dependent) for either queen phenotype, but colonies at both sites produced numerous gynes, and we infer that all foundresses initiate colonies independently and are obligate foragers.