Heteroponera

Most specimens reviewed here were collected as foraging workers taken in Berlese funnel samples, siftings of forest leafmould or moss, or stick brushings. Colonies of Heteroponera relicta, Heteroponera rhodopygea and Heteroponera lioprocta are found in and under rotting wood fragments or logs on the forest floor, or in soil under stones (pers obs.). Little is known of other species. Living H. relicta and H. rhodopygea frequently carry numbers of very small phoretic mites clustered in their antennal scrobes or elsewhere. Their remains may be present on cabinet specimens, including those of other species. Their systematics and phylogeny relative to the host ants would likely be of scientific interest. Heteroponera workers roll their bodies ventrally when disturbed, with the antennae enclosed in the scrobes, and are sometimes almost imperceptible against the substrate in this position. The structure of the head/ mesosoma interface including the pronotal epaulets, and the gastral configuration are likely adapted to this purpose. (Taylor 2015, for northeastern Australia Heteroponera relicta group species).

Identification
Taylor (2015) - Heteroponera may be identified in Australia using the keys of Hoelldobler & Wilson (1990), Bolton (1994) and Shattuck (1999), notably regarding distinction from the ectatommine genus Rhytidoponera. Note however that some small sympatric Rhytidoponera species (relatives of Rhytidoponera tenuis Forel —see Brown, 1958, Appendix: notes 15, 16, 27.) commonly present in North Queensland berlesates and other bulk samples which include H. relicta-group specimens, lack allegedly diagnostic supplementary teeth on the pretarsal claws, but are otherwise readily discriminated from Heteroponera.

Heteroponera species groups

Distribution
Taylor (2015) - Heteroponera species are known only from Australia, New Zealand, and the Neotropics, with putative southern, Gondwanic, biogeographic connections linking the amphipacific faunas (Feitosa, 2011; Taylor, 2011.

Species by Region
Number of species within biogeographic regions, along with the total number of species for each region.

Biology
Taylor (2015) - The nine new species described in the formerly monotypic species-group of Heteroponera relicta (Wheeler, 1915) raise the tally of named Australasian Heteroponera species to sixteen (Australia 15: New Zealand 1). The H. relicta-group, in which two species complexes are recognized, is known only from rainforest localities centered on the Wet Tropics World Heritage Area of Northeast Queensland. Its remarkably concentrated overall distributional range spans approximately 6 degrees of latitude from near Cooktown (15°28'S, 145°15'E) south to the vicinity of Cannonvale (20°16’S, 148°43’E), a latitudinal distance of only ca. 550 km. in which no collection record is more than ca. 35 km from the coast. This area covers less than 1% of the Australian continent, but supports a major proportion of its biota. The H. relicta group is thus regionally endemic in the sense of Yeates et al. (2002).

Taylor (2011) recognized three Australasian species groups in Heteroponera following Brown (1958), and reviewed the H. leae group (H. leae (Wheeler), H. crozieri Taylor and H. majeri Taylor). The remaining group of H. imbellis (Emery) includes the New Zealand endemic H. brouni (Forel) and several Australian nominal species listed as synonyms of H. imbellis by Brown (1958).

Feitosa (pers comm.) currently recognizes 14 Neotropical species (13 formally named, two probable junior synonyms, and three undescribed species).

Taxonomic distinction of the relicta, leae and imbellis groups in Australia is supported by Feitosa’s (2011) phylogenetic analysis. In his cladograms (ibid., fig 4): (1) The leae-group species H. leae and H. crozieri (the latter given as “Heteroponera sp B”) are sister taxa and share a common higher-order clade with H. brouni and three Neotropical species (Interestingly, those taxa: H. inca Brown, H. monticola Kempf & Brown, H. carinifrons and another undescribed species, are the only known Neotropical Heteroponera found exclusively west of the Andes, and the South Chilean H. carinifrons, a sister to the New Zealand H. brouni in Feitosa’s analysis, is the most southern known Neotropical Heteroponera. (2) That higher-order clade is sister to one which includes the two analyzed Australian imbellis-group species (cited as “H. imbellis” and “H. sp A”), along with 11 further Neotropical species. (3) The clade including all of the above taxa is sister to another which comprises the two analyzed H. relicta–group species, H. relicta and H. rhodopygea (the latter as “Heteroponera sp C”). The H. relicta-group, reviewed here, is thus represented as a sister lineage to the clade including all other Heteroponera species analyzed by Feitosa (whether Australasian or Neotropical). Its constituent species are thus arguably relictual on the Australian continent as the least morphologically–derived of all Heteroponera species now extant.

Feitosa’s findings provide compelling evidence that major lineages in Heteroponera, destined eventually to include both Australasian and Neotropical species, were almost certainly represented on Gondwana prior to the Jurassic separation of the Australian and South American continental plates. Phylogenetically the two modern amphipacific faunas are geographically separated derivatives of a single ancient Gondwanic radiation. Heteroponera was thus evidentially substantially diversified on Gondwana, doubtless including species very like those existing today, at least ca 150 million years ago, and the genus must have originated before that time, presumably on Gondwana.

Castes
A few species have ergatoid queens (flightless) (Heteroponera brounii, Heteroponera carinifrons, Heteroponera inca, Heteroponera mayri, Heteroponera monticola, Heteroponera relicta). This is likely to reflect a shift to Dependent Colony Foundation [Life History]. Ergatoid queens can function as secondary reproductives (i.e. they succeed a winged foundress) in some species.

Nomenclature

 *  HETEROPONERA [Heteroponerinae]
 * Heteroponera Mayr, 1887: 532. Type-species: Heteroponera carinifrons, by monotypy.
 * Heteroponera junior synonym of Acanthoponera: Emery, 1911d: 35; Wheeler, W.M. 1922a: 643; Wheeler, 1923e: 179.
 * Heteroponera subgenus of Acanthoponera: Forel, 1917: 236.
 * Heteroponera revived status as genus: Brown, 1952h: 70.
 * Heteroponera senior synonym of Anacanthoponera: Brown, 1952h: 70; Brown, 1958g: 194.
 * Heteroponera senior synonym of Paranomopone: Brown, in Borgmeier, 1957: 112; Brown, 1958g: 194.
 * ANACANTHOPONERA [junior synonym of Heteroponera]
 * Anacanthoponera Wheeler, W.M. 1923e: 176 [as subgenus of Acanthoponera]. Type-species: Ponera dolo, by original designation.
 * Anacanthoponera junior synonym of Heteroponera: Brown, 1952h: 70.
 * PARANOMOPONE [junior synonym of Heteroponera]
 * Paranomopone Wheeler, W.M. 1915c: 117. Type-species: Paranomopone relicta, by monotypy.
 * Paranomopone junior synonym of Heteroponera: Brown, in Borgmeier, 1957: 112; Brown, 1958g: 194.