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Abstract The classification of ants (Hymenoptera: Formicidae) has progressed in waves since the first 17 species were described by Linnaeus in the 1758 edition of Systema Naturae. Since then, over 18,000 species-rank names have accumulated for the global myrmecofauna, of which ~14,260 living and ~810 fossil species are valid. Here, we provide a synopsis of ant biodiversity and review the history and classification of the family, while highlighting the massive growth of the field in the new millennium. We observe that major transformation has occurred for ant classification due to advances in DNA sequencing technologies, model-based hypothesis testing, and imaging technologies. We therefore provide a revised and illustrated list of diagnostic character states for the higher clades of Formicidae, recognizing that vastly more work is to be done. To facilitate discussion and the systematic accumulation of evolutionary knowledge for the early evolution of the ants, we suggest an informal nomenclatural system for the higher clades of ants, based on names currently in use and a set of names that have been democratically selected by the authors. To guide future work on ant systematics, we summarize currently available databases and present perspectives on regions in need of biodiversity exploration, challenges facing the field, and the future of ant taxonomy. 

Fungus-farming ants cultivate multiple lineages of fungi for food, but, because fungal cultivar relationships are largely unresolved, the history of fungus-ant coevolution remains poorly known. We designed probes targeting >2000 gene regions to generate a dated evolutionary tree for 475 fungi and combined it with a similarly generated tree for 276 ants. We found that fungus-ant agriculture originated ~66 million years ago when the end-of-Cretaceous asteroid impact temporarily interrupted photosynthesis, causing global mass extinctions but favoring the proliferation of fungi. Subsequently, ~27 million years ago, one ancestral fungal cultivar population became domesticated, i.e., obligately mutualistic, when seasonally dry habitats expanded in South America, likely isolating the cultivar population from its free-living, wet forest–dwelling conspecifics. By revealing these and other major transitions in fungus-ant coevolution, our results clarify the historical processes that shaped a model system for nonhuman agriculture. 

Abstract The evolutionary history of fungus-farming ants has been the subject of multiple morphological, molecular phylogenetic, and phylogenomic studies. Due to its rarity, however, the phylogenetic position, natural history, and fungal associations of the monotypic genus Paramycetophylax Kusnezov have remained enigmatic. Here we report the first excavations of colonies of Paramycetophylax bruchi (Santschi) and describe its nest architecture and natural history. Utilizing specimens from these collections, we generated ultraconserved-element (UCE) data to determine the evolutionary position of Paramycetophylax within the fungus-farming ants and ribosomal ‘fungal barcoding’ ITS sequence data to identify the fungal cultivar. A maximum-likelihood phylogenomic analysis indicates that the genus Paramycetophylax is the sister group of the yeast-cultivating Cyphomyrmex rimosus group, an unexpected result that renders the genus Cyphomyrmex Mayr paraphyletic. A Bayesian divergence-dating analysis indicates that Paramycetophylax diverged from its sister group around 36 mya (30–42 mya, HPD) in the late Eocene-early Oligocene, a period of global cooling, expansion of grasslands, and large-scale extinction of tropical organisms. Bayesian analysis of the fungal cultivar ITS gene fragment indicates that P. bruchi practices lower agriculture and that the cultivar grown by P. bruchi belongs to the Clade 1 group of lower-attine fungi, a clade that, interestingly, also includes the C. rimosus-group yeast cultivars. Based on these results, we conclude that a better understanding of P. bruchi and its fungal cultivar, including whole-genome data, is critical for reconstructing the origin of yeast agriculture, a major transition in the evolution of fungus-farming ants. 

Fungi shape the diversity of life. Characterizing the evolution of fungi is critical to understanding symbiotic associations across kingdoms. In this study, we investigate the genomic and metabolomic diversity of the genus Escovopsis , a specialized parasite of fungus-growing ant gardens. Based on 25 high-quality draft genomes, we show that Escovopsis forms a monophyletic group arising from a mycoparasitic fungal ancestor 61.82 million years ago (Mya). Across the evolutionary history of fungus-growing ants, the dates of origin of most clades of Escovopsis correspond to the dates of origin of the fungus-growing ants whose gardens they parasitize. We reveal that genome reduction, determined by both genomic sequencing and flow cytometry, is a consistent feature across the genus Escovopsis, largely occurring in coding regions, specifically in the form of gene loss and reductions in copy numbers of genes. All functional gene categories have reduced copy numbers, but resistance and virulence genes maintain functional diversity. Biosynthetic gene clusters (BGCs) contribute to phylogenetic differences among Escovopsis spp., and sister taxa in the Hypocreaceae. The phylogenetic patterns of co-diversification among BGCs are similarly exhibited across mass spectrometry analyses of the metabolomes of Escovopsis and their sister taxa. Taken together, our results indicate that Escovopsis spp. evolved unique genomic repertoires to specialize on the fungus-growing ant-microbe symbiosis. 

Abstract Although calcareous anatomical structures have evolved in diverse animal groups, such structures have been unknown in insects. Here, we report the discovery of high-magnesium calcite [CaMg(CO 3 ) 2 ] armor overlaying the exoskeletons of major workers of the leaf-cutter ant Acromyrmex echinatior . Live-rearing and in vitro synthesis experiments indicate that the biomineral layer accumulates rapidly as ant workers mature, that the layer is continuously distributed, covering nearly the entire integument, and that the ant epicuticle catalyzes biomineral nucleation and growth. In situ nanoindentation demonstrates that the biomineral layer significantly hardens the exoskeleton. Increased survival of ant workers with biomineralized exoskeletons during aggressive encounters with other ants and reduced infection by entomopathogenic fungi demonstrate the protective role of the biomineral layer. The discovery of biogenic high-magnesium calcite in the relatively well-studied leaf-cutting ants suggests that calcareous biominerals enriched in magnesium may be more common in metazoans than previously recognized. 

Evolutionary adaptations for maintaining beneficial microbes are hallmarks of mutualistic evolution. Fungus-farming “attine” ant species have complex cuticular modifications and specialized glands that house and nourish antibiotic-producing Actinobacteria symbionts, which in turn protect their hosts’ fungus gardens from pathogens. Here we reconstruct ant–Actinobacteria evolutionary history across the full range of variation within subtribe Attina by combining dated phylogenomic and ultramorphological analyses. Ancestral-state analyses indicate the ant–Actinobacteria symbiosis arose early in attine-ant evolution, a conclusion consistent with direct observations of Actinobacteria on fossil ants in Oligo-Miocene amber. qPCR indicates that the dominant ant-associated Actinobacteria belong to the genus Pseudonocardia . Tracing the evolutionary trajectories of Pseudonocardia -maintaining mechanisms across attine ants reveals a continuum of adaptations. In Myrmicocrypta species, which retain many ancestral morphological and behavioral traits, Pseudonocardia occur in specific locations on the legs and antennae, unassociated with any specialized structures. In contrast, specialized cuticular structures, including crypts and tubercles, evolved at least three times in derived attine-ant lineages. Conspicuous caste differences in Pseudonocardia -maintaining structures, in which specialized structures are present in worker ants and queens but reduced or lost in males, are consistent with vertical Pseudonocardia transmission. Although the majority of attine ants are associated with Pseudonocardia , there have been multiple losses of bacterial symbionts and bacteria-maintaining structures in different lineages over evolutionary time. The early origin of ant– Pseudonocardia mutualism and the multiple evolutionary convergences on strikingly similar anatomical adaptations for maintaining bacterial symbionts indicate that Pseudonocardia have played a critical role in the evolution of ant fungiculture.