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Despite recent taxonomically focused molecular phylogenies [37,38,39] and extensive taxonomic works [12, 31], Temnothorax remains widely renowned as one of the most taxonomically difficult ant genera [40]: its status as a phylogenetically natural group has been repeatedly revised [41] and called into question [42]. This is a product of multiple factors: subtle morphological variation on a global scale among groups of species, striking departures from the morphological norm in others, morphological convergence with distantly related genera [2], weak diagnostic characters for the genus as a whole, and regional treatments of species in the absence of global understanding of the clade structure (see Additional file 1 for a more detailed account of recent molecular phylogenetic work and the taxonomic history of Temnothorax). The current absence of an infra-generic classification system for Temnothorax is not for a lack of effort, though: past taxonomists erected five subgenera to compartmentalize the species diversity of this genus, none of which are currently recognized due to a lack of clear diagnostic characters [41]. In this study, I estimate the phylogeny of Temnothorax on a global scale to gain insight into the structure of relationships within the genus and, furthermore, to verify that the genus is monophyletic as currently diagnosed. To maximize within-genus lineage diversity, I sampled broadly from the species groups that have been formally designated in the literature. I simultaneously examine several key aspects of the historical biogeography and evolution in the genus, which I present below.
The phylogenetic analyses from two sequencing approaches and on multiple data subsets recover identical clades in Temnothorax and largely agree on the relationships among them. Although even the large UCE dataset could not resolve some of the relationships in the backbone satisfactorily, constraint analyses on the Sanger dataset resolved most of the difficult nodes. Temnothorax was recovered as a natural group with high support, as was the sister group relationship the Leptothorax genus group. The species groups, former subgenera, and satellite genera of the core formicoxenines, including Temnothorax have a long and controversial history. A phylogenetic framework for understanding the relationships within this intriguing group of ants has been sorely lacking, which has hampered taxonomic revisionary efforts and our understanding of the pace and mode of evolution within this genus. In this study, the inclusion of the former genera Chalepoxenus, Myrmoxenus, and Protomognathus in Temnothorax is found to be justified. The former genus Macromischa is revealed to be polyphyletic, confirming earlier suppositions [12, 41], and composed of members of the salvini, obturator, and sallei clades, interdigitated with the former subgenus Myrafant. A striking pattern within Temnothorax is strong morphological conservatism within and between groups (e.g., compare taxa from the tricarinatus group and the andrei, rugatulus, and Palearctic clades) punctuated by extreme morphological departures (e.g. the Caribbean members of the sallei clade, and the salvini clade). 2ff7e9595c
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