The evolution of myrmicine ants: phylogeny and biogeography of a hyperdiverse ant clade (<scp>H</scp>ymenoptera:<scp>F</scp>ormicidae)

  • PHILIP S. WARD
    Department of Entomology and Nematology University of California Davis CA U.S.A.
  • SEÁN G. BRADY
    Department of Entomology National Museum of Natural History, Smithsonian Institution Washington DC U.S.A.
  • BRIAN L. FISHER
    Department of Entomology California Academy of Sciences San Francisco CA U.S.A.
  • TED R. SCHULTZ
    Department of Entomology National Museum of Natural History, Smithsonian Institution Washington DC U.S.A.

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<jats:title>Abstract</jats:title><jats:p>This study investigates the evolutionary history of a hyperdiverse clade, the ant subfamily<jats:styled-content style="fixed-case">M</jats:styled-content>yrmicinae (<jats:styled-content style="fixed-case">H</jats:styled-content>ymenoptera:<jats:styled-content style="fixed-case">F</jats:styled-content>ormicidae), based on analyses of a data matrix comprising 251 species and 11 nuclear gene fragments. Under both maximum likelihood and<jats:styled-content style="fixed-case">B</jats:styled-content>ayesian methods of inference, we recover a robust phylogeny that reveals six major clades of<jats:styled-content style="fixed-case">M</jats:styled-content>yrmicinae, here treated as newly defined tribes and occurring as a pectinate series:<jats:styled-content style="fixed-case">M</jats:styled-content>yrmicini,<jats:styled-content style="fixed-case">P</jats:styled-content>ogonomyrmecini<jats:bold>trib.n.</jats:bold>,<jats:styled-content style="fixed-case">S</jats:styled-content>tenammini,<jats:styled-content style="fixed-case">S</jats:styled-content>olenopsidini,<jats:styled-content style="fixed-case">A</jats:styled-content>ttini and<jats:styled-content style="fixed-case">C</jats:styled-content>rematogastrini. Because we condense the former 25 myrmicine tribes into a new six‐tribe scheme, membership in some tribes is now notably different, especially regarding<jats:styled-content style="fixed-case">A</jats:styled-content>ttini. We demonstrate that the monotypic genus<jats:italic>Ankylomyrma</jats:italic>is neither in the<jats:styled-content style="fixed-case">M</jats:styled-content>yrmicinae nor even a member of the more inclusive formicoid clade—rather it is a poneroid ant, sister to the genus<jats:italic>Tatuidris</jats:italic>(<jats:styled-content style="fixed-case">A</jats:styled-content>groecomyrmecinae). Several species‐rich myrmicine genera are shown to be nonmonophyletic, including<jats:italic>Pogonomyrmex</jats:italic>,<jats:italic>Aphaenogaster</jats:italic>,<jats:italic>Messor</jats:italic>,<jats:italic>Monomorium</jats:italic>,<jats:italic>Pheidole</jats:italic>,<jats:italic>Temnothorax</jats:italic>and<jats:italic>Tetramorium</jats:italic>. We propose a number of generic synonymies to partially alleviate these problems (senior synonym listed first):<jats:italic>Pheidole</jats:italic>=<jats:italic>Anisopheidole</jats:italic><jats:bold>syn.n.</jats:bold> = <jats:italic>Machomyrma</jats:italic><jats:bold>syn.n.</jats:bold>;<jats:italic>Temnothorax</jats:italic>=<jats:italic>Chalepoxenus</jats:italic><jats:bold>syn.n.</jats:bold> = <jats:italic>Myrmoxenus</jats:italic><jats:bold>syn.n.</jats:bold> = <jats:italic>Protomognathus</jats:italic><jats:bold>syn.n.</jats:bold>;<jats:italic>Tetramorium</jats:italic>=<jats:italic>Rhoptromyrmex</jats:italic><jats:bold>syn.n.</jats:bold> = <jats:italic>Anergates</jats:italic><jats:bold>syn.n.</jats:bold>=<jats:italic>Teleutomyrmex</jats:italic><jats:bold>syn.n.</jats:bold>The genus<jats:italic>Veromessor</jats:italic><jats:bold>stat.r.</jats:bold>is resurrected for the<jats:styled-content style="fixed-case">N</jats:styled-content>ew<jats:styled-content style="fixed-case">W</jats:styled-content>orld species previously placed in<jats:italic>Messor</jats:italic>;<jats:italic>Syllophopsis</jats:italic><jats:bold>stat.r.</jats:bold>is resurrected from synonymy under<jats:italic>Monomorium</jats:italic>to contain the species in the<jats:italic>hildebrandti</jats:italic>group;<jats:italic>Trichomyrmex</jats:italic><jats:bold>stat.r.</jats:bold>is resurrected from synonymy under<jats:italic>Monomorium</jats:italic>to contain the species in the<jats:italic>scabriceps</jats:italic>‐ and<jats:italic>destructor</jats:italic>‐groups; and the monotypic genus<jats:italic>Epelysidris</jats:italic><jats:bold>stat.r.</jats:bold>is reinstated for<jats:italic>Monomorium brocha</jats:italic>. Bayesian divergence dating indicates that the crown group<jats:styled-content style="fixed-case">M</jats:styled-content>yrmicinae originated about 98.6 Ma (95% highest probability density 87.9–109.6 Ma) but the six major clades are considerably younger, with age estimates ranging from 52.3 to 71.1 Ma. Although these and other suprageneric taxa arose mostly in the middle<jats:styled-content style="fixed-case">E</jats:styled-content>ocene or earlier, a number of prominent, species‐rich genera, such as<jats:italic>Pheidole</jats:italic>,<jats:italic>Cephalotes</jats:italic>,<jats:italic>Strumigenys</jats:italic>,<jats:italic>Crematogaster</jats:italic>and<jats:italic>Tetramorium</jats:italic>, have estimated crown group origins in the late<jats:styled-content style="fixed-case">E</jats:styled-content>ocene or<jats:styled-content style="fixed-case">O</jats:styled-content>ligocene. Most myrmicine species diversity resides in the two sister clades,<jats:styled-content style="fixed-case">A</jats:styled-content>ttini and<jats:styled-content style="fixed-case">C</jats:styled-content>rematogastrini, which are estimated to have originated and diversified extensively in the<jats:styled-content style="fixed-case">N</jats:styled-content>eotropics and<jats:styled-content style="fixed-case">P</jats:styled-content>aleotropics, respectively. The newly circumscribed<jats:styled-content style="fixed-case">M</jats:styled-content>yrmicini is<jats:styled-content style="fixed-case">H</jats:styled-content>olarctic in distribution, and ancestral range estimation suggests a<jats:styled-content style="fixed-case">N</jats:styled-content>earctic origin. The<jats:styled-content style="fixed-case">P</jats:styled-content>ogonomyrmecini and<jats:styled-content style="fixed-case">S</jats:styled-content>olenopsidini are reconstructed as being<jats:styled-content style="fixed-case">N</jats:styled-content>eotropical in origin, but they have subsequently colonized the<jats:styled-content style="fixed-case">N</jats:styled-content>earctic region (<jats:styled-content style="fixed-case">P</jats:styled-content>ogonomyrmecini) and many parts of the<jats:styled-content style="fixed-case">O</jats:styled-content>ld<jats:styled-content style="fixed-case">W</jats:styled-content>orld as well as the<jats:styled-content style="fixed-case">N</jats:styled-content>earctic region (<jats:styled-content style="fixed-case">S</jats:styled-content>olenopsidini), respectively. The<jats:styled-content style="fixed-case">S</jats:styled-content>tenammini have flourished primarily in the northern hemisphere, and are most likely of<jats:styled-content style="fixed-case">N</jats:styled-content>earctic origin, but selected lineages have dispersed to the northern<jats:styled-content style="fixed-case">N</jats:styled-content>eotropics and the<jats:styled-content style="fixed-case">P</jats:styled-content>aleotropics. Thus the evolutionary history of the<jats:styled-content style="fixed-case">M</jats:styled-content>yrmicinae has played out on a global stage over the last 100 Ma, with no single region being the principal generator of species diversity.</jats:p><jats:p>This published work has been registered in ZooBank,<jats:ext-link xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="http://zoobank.org/urn:lsid:zoobank.org:pub:%20BB6829C4-DA79-45FE-979E-9749E237590E">http://zoobank.org/urn:lsid:zoobank.org:pub: BB6829C4‐DA79‐45FE‐979E‐9749E237590E</jats:ext-link>.</jats:p>

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