Phylogenetics and Biogeography of Ponerine Ants Chris A. Schmidt

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Phylogenetics and Biogeography of Ponerine Ants

Chris A. Schmidt

Ponerine ants (Ponerinae) are a diverse and conspicuous component of tropical faunas. With
nearly 1,000 described species worldwide they constitute the third largest ant subfamily, and as a
group ponerines exhibit impressive morphological, ecological, and behavioral diversity. Despite
their ecological importance and value as potential models for studying social evolution, the
Ponerinae has never recieved any phylogenetic treatment, and the relationships among the major
lineages and their biogeographic history are almost completely unknown. Major systematic
challenges remain, as past reliance on morphological evidence and phenetic (rather than
cladistic) principles has resulted in an untenable status quo in ponerine taxonomy.
I present a preliminary molecular phylogeny for the Ponerinae, utilizing nuclear and
mitochondrial protein-coding genes and nuclear ribosomal genes to elucidae tribal and generic
relationships. The tribes Platythyreini and Ponerini are found as sister groups with high statistical
support (the third tribe, Thaumatomyrmecini, is exceedingly rare and has not yet been sampled).
The large genus Pachycondyla is found to be massively polyphyletic, with members scattered
throughout the Ponerini, illustrating the need for a revision of ponerine taxonomy at the genus
level. A tentative redefinition of ponerine genera is given, with nearly 40 genera recognized
(compared with 25 currently recognized genera). At the broadest levels, four major clades are
recognized within Ponerini: the Southeast Asian genus Diacamma; a large clade containing all
hypogeic (cryptic foraging) ponerine taxa sampled, including Hypoponera; a diverse Neotropical
clade including Pachycondyla sensu stricto; and a large Old World clade of epigeic (non-cryptic
foraging) ponerines which includes Leptogenys, Odontomachus, and many other groups. The
poor resolution and long branches within this latter Old World clade suggest that it underwent a
rapid and rich radiation deep in the past.
Results of a preliminary phylogenetic dating analysis are presented, with a discussion of
its implications for ponerine biogeography. The two major ponerine tribes are estimated to have
diverged roughly 110 million years ago, much older than the splits between some long-
recognized ant subfamilies, suggesting that Platythyreini should perhaps be raised to subfamily
status. Relationships among other ant subfamilies (widely sampled as outgroups) are also briefly
discussed. I find support for Leptanillinae as the sister group to all other extant ants, with an
unresolved grade of “poneroid” taxa (Proceratiinae, Amblyoponinae, and Paraponerinae), and
finally Ponerinae as sister to the remaining ant subfamilies which together constitute roughly
85% of the described ant species diversity.
Among other results, mass raiding (i.e. army ant-like foraging behavior) is shown to have
arisen independently at least 4 times among the Ponerini. The deep phylogenetic split between
hypogeic and epigeic ponerines is an unexpected result and contradicts the previous belief that
hypogeic ponerines arose multiple times from epigeic ancestors. Another result is that
reproduction by gamergates (mated, reproductive workers) has arisen multiple times within the
Ponerini. Gamergates are rare outside the Ponerini, and their frequent evolution within the
Ponerini highlights the extensive diversity of ponerine social systems.
These results should ultimately help reconstruct the sequence of evolutionary, migratory,
and geological events that resulted in the present diversity and distribution of the subfamily, and
will lay the groundwork for future taxonomic revisions and studies of ponerine morphological,
ecological, and social evolution.

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