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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. 

Abstract The spectacular radiation of insects has produced a stunning diversity of phenotypes. During the past 250 years, research on insect systematics has generated hundreds of terms for naming and comparing them. In its current form, this terminological diversity is presented in natural language and lacks formalization, which prohibits computer-assisted comparison using semantic web technologies. Here we propose a Model for Describing Cuticular Anatomical Structures (MoDCAS) which incorporates structural properties and positional relationships for standardized, consistent, and reproducible descriptions of arthropod phenotypes. We applied the MoDCAS framework in creating the ontology for the Anatomy of the Insect Skeleto-Muscular system (AISM). The AISM is the first general insect ontology that aims to cover all taxa by providing generalized, fully logical, and queryable, definitions for each term. It was built using the Ontology Development Kit (ODK), which maximizes interoperability with Uberon (Uberon multi-species anatomy ontology) and other basic ontologies, enhancing the integration of insect anatomy into the broader biological sciences. A template system for adding new terms, extending, and linking the AISM to additional anatomical, phenotypic, genetic, and chemical ontologies is also introduced. The AISM is proposed as the backbone for taxon-specific insect ontologies and has potential applications spanning systematic biology and biodiversity informatics, allowing users to (1) use controlled vocabularies and create semi-automated computer-parsable insect morphological descriptions; (2) integrate insect morphology into broader fields of research, including ontology-informed phylogenetic methods, logical homology hypothesis testing, evo-devo studies, and genotype to phenotype mapping; and (3) automate the extraction of morphological data from the literature, enabling the generation of large-scale phenomic data, by facilitating the production and testing of informatic tools able to extract, link, annotate, and process morphological data. This descriptive model and its ontological applications will allow for clear and semantically interoperable integration of arthropod phenotypes in biodiversity studies. 

Camponotus and Colobopsis are widely distributed and species-rich genera in the ant tribe Camponotini. Molecular phylogenetic studies demonstrate that they are not sister taxa, but several lineages within each genus have converged to a remarkable degree, confounding the taxonomy of these ants. Based on multiple lines of evidence, including worker and male morphology, we demonstrate that: (1) three species of “Camponotus” belonging to the subgenus Myrmotemnus, including its type species, are in fact members of the genus Colobopsis ; (2) four species previously assigned to Colobopsis belong to the subgenus Myrmamblys of Camponotus ; and (3) three Nearctic taxa recently placed in Colobopsis are members of the genus Camponotus and closely related to Camponotus clarithorax . These taxonomic findings yield the following new or revived combinations: Colobopsis moeschi ( comb. nov. ), Colobopsis moeschi lygaea ( comb. nov. ), Colobopsis nutans ( comb. nov. ), Colobopsis nutans cleliae ( comb. nov. ), and Colobopsis reichenspergeri ( comb. nov. ); Camponotus apostemata ( comb. nov. ), Camponotus aurelianus ( comb. rev. ), Camponotus cavibregma ( comb. nov. ), Camponotus horrens ( comb. rev. ), Camponotus politae ( comb. rev. ), Camponotus trajanus ( comb. rev. ), and Camponotus yogi ( comb. rev. ). A further consequence is the following generic synonymy (senior synonym listed first): Colobopsis = Myrmotemnus syn. nov. , and Camponotus = Dolophra syn. rev. At the species level, we argue that Camponotus apostemata and Camponotus cavibregma are junior synonyms ( syn. nov. ) of Camponotus yogi , and Camponotus quercicola is a junior synonym ( syn. nov. ) of Ca. laevigatus . Taxonomic comments are also provided on some members of the Camponotus reticulatus group, with Camponotus adustus ( stat. nov. ) and Ca. leucodiscus ( stat. rev. ) being recognized as distinct species rather than subspecies of Ca. bellus . A male-based diagnosis of the Camponotini is provided, and differences between the males of Colobopsis and Camponotus are documented and illustrated for the first time. This study reveals new character systems of potential value to the systematics of these ants, including features of the male genitalia, and emphasizes the value of reciprocal illumination between phylogenomics and critical morphological analysis.