Schmidt, Chris, 2013, Molecular phylogenetics of ponerine ants (Hymenoptera: Formicidae: Ponerinae), Zootaxa 3647 (2), pp. 201-250: 209-212

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Monophyly of Ponerinae  and its tribes and genera

The monophyly of Ponerinae  was strongly supported by analyses of the ALL_DATA matrix (BPP = 1.00, BS = 0.98; Fig. 3 View Figure ), confirming the results of previous molecular phylogenetic studies with more limited sampling of ponerine taxa (e.g., Moreau et al., 2006; Brady et al., 2006) and confirming predictions from morphological evidence. In his morphological synopsis of Ponerinae  , Bolton (2003) noted two synapomorphies for the subfamily: fusion of the toruli to the frontal lobes, and the shape of the outer borders of the frontal lobes, which form ”simple short semicircles or blunt triangles and in full-face view have a distinctly pinched-in appearance posteriorly”. An examination of the inferred phylogeny of Formicidae  and the subfamily-level morphological synopses of Bolton (2003) suggest an additional possible synapomorphy for Ponerinae  , the loss of tergosternal fusion of the petiole (C. Schmidt and S. Shattuck, in prep), though this trait could conceivably be symplesiomorphic, with repeated evolution of a fused petiole in other poneroid lineages. All current ponerine genera not sampled in this study ( Asphinctopone  , Belonopelta  , Boloponera  , Dolioponera  , Feroponera  , and Promyopias  ) display these traits and their placements within Ponerinae  and Ponerini  are not in doubt, so it is highly unlikely that their inclusion in this study would have rendered Ponerinae  non-monophyletic.

Phylogenetic analysis of the ALL_DATA matrix ( Fig. 3 View Figure ) strongly supported the monophyly of Platythyreini  ( Platythyrea  ) (BPP = 1.00, BS = 1.00). The status of the genus Eubothroponera  as a junior synonym of Platythyrea  (Brown, 1952, 1975) is here confirmed with molecular data for the first time: P. t u r n e r i, though never formally included in Eubothroponera  despite its obvious close relationship to members of that former genus, is nested within Platythyrea  with strong support (monophyly of Platythyrea  excluding P. t u r n e r i: BPP <0.0001; BS = 0.002). Members of Eubothroponera  have several morphological traits that differ from most other Platythyrea  species, including coarser sculpturing, denser standing pilosity but sparser pubescence, and a relatively low helcium. The latter character approximates the condition found in most Ponerini  , but the results of this study imply that low helcia were independently derived in Platythyrea  and Ponerini  .

The monophyly of the diverse tribe Ponerini  was not supported in either Bayesian or ML bootstrap analyses (BPP <0.0001; BS <0.001), though Ponerini  monophyly could not be rejected by an SH test (p -value = 0.661). The non-monophyly of Ponerini  stems from its inclusion of Thaumatomyrmecini  ( Thaumatomyrmex  ), a result also recovered by Brady et al. (2006). The phylogenetic position of Thaumatomyrmex  has been unclear due to its bizarre structure (including pitchfork-like mandibles, widely separated frontal lobes, and unusually convex eyes; Kempf, 1975). Bolton (2003) noted that Thaumatomyrmex  shares several apomorphies with Ponerini  , including vestigial male mandibles and articulation of the petiole low on the anterior face of the first gastral segment, but maintained its separate tribal status. The genus lacks a principal synapomorphy of Ponerini  (the narrow insertion of the clypeus between the frontal lobes, and related characters; Bolton, 2003), but this character is highly modified in Thaumatomyrmex  , rendering it uninformative about the phylogenetic position of the genus. Morphological data therefore imply at least a sister relationship between Thaumatomyrmex  and Ponerini  , and are consistent with its placement within Ponerini  . The molecular results reported here suggest that Thaumatomyrmex  is simply a highly derived member of Ponerini  , its strange morphological traits secondarily derived as a result of extreme prey specialization (Brandão, 1991).

Among sequenced taxa, all sampled ponerine genera were strongly inferred to be monophyletic (BPP and BS = 1.00 for all genera except Myopias  , for which BPP and BS both = 0.99, and Thaumatomyrmex  , for which BS = 0.99) except Pachycondyla  and Odontomachus  . Pachycondyla  was strongly inferred to be non-monophyletic (BPP of monophyletic Pachycondyla  <0.0001; BS <0.001; SH p- value <0.00001), as members of this genus were broadly distributed across the phylogeny of Ponerini  . The sampled Pachycondyla  species represent numerous distinct lineages which largely correspond to the boundaries of Pachycondyla  “subgenera,” though Bothroponera  , Mesoponera  and Pseudoponera  are themselves non-monophyletic ( Bothroponera  : BPP <0.001, BS = 0.014, SH p - value = 0.768; Mesoponera  : BPP <0.0001, BS <0.001, SH p -value <0.00001; Pseudoponera  : BPP <0.0001, BS <0.001, SH p -value <0.001).

Pachycondyla  is the end product of heavy synonymization at the genus level, due primarily to the work of W. L. Brown, who was in the process of revising ponerine taxonomy at the time of his death. Brown’s formal revision remains unpublished, but his numerous planned synonymizations have been published elsewhere without supporting justification (Brown, 1973; Snelling, 1981; Hölldobler and Wilson, 1990; Bolton, 1994; see also Mackay and Mackay, 2010) and in practice have been generally accepted by the scientific community. The monophyly of Pachycondyla  is doubtful even in the absence of molecular data because the genus displays substantial morphological, ecological and behavioral diversity (reviewed in C. Schmidt and S. Shattuck, in prep.), is defined only by morphological symplesiomorphies such as triangular mandibles and paired metatibial spurs, and contains several lineages that show clear morphological similarities to other ponerine genera. This study confirms the extensive non-monophyly of Pachycondyla  and stresses the need for a comprehensive taxonomic revision of ponerine genera (provided in C. Schmidt and S. Shattuck, in prep.).

Analysis of the ALL_DATA matrix was ambiguous about Odontomachus  monophyly, with three resolutions: O. coquereli Roger  sister to Anochetus  (BPP = 0.58, BS = 0.37), Odontomachus  monophyletic (BPP = 0.29, BS = 0.41), or O. coquereli  sister to Anochetus  + other Odontomachus  (BPP = 0.13, BS = 0.22). To further address this question, a preliminary expanded data set was produced which included sequences for six additional Anochetus  taxa and 18 additional Odontomachus  taxa, plus data for the mitochondrial gene cytochrome oxidase I (COI), but excluding abdA, EF 1 αF 1, and EF 1 αF 2 (C. Schmidt, unpublished data). Phylogenetic analyses of these data supported reciprocal monophyly of Odontomachus  and Anochetus  (results not shown), but some phylogenetically important Anochetus  species (such as members of the A. gladiator (Mayr)  and A. cato Forel  species groups; Brown, 1976, 1978) were not sampled and their inclusion in future phylogenetic analyses could still reveal the nonmonophyly of either genus.