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Redefinition of the Amblyoponini HNS
Long considered a rather primitive group of ants, possibly reflective of an early stage in ant evolution (Wilson, 1971; Hoelldobler & Wilson, 1990), the Amblyoponini HNS were well characterized by Brown (1960; see also Brown, 1974 a, b; Gotwald & Levieux, 1972) but they have not heretofore been defined explicitly in terms of derived characters that would signify their monophyly. The characterization given below refers to all currently recognized, extant amblyoponine genera for which the worker caste is known (i. e. Amblyopone HNS , Concoctio HNS , Mystrium HNS , Myopopone HNS , Onychomyrmex HNS and Prionopelta HNS ). These were compared to a representative range of taxa in other ponerine tribes ( Ectatommini HNS . Platythyreini HNS , Ponerini HNS , Typhlomyrmecini HNS ) as well as to Apomyrma HNS and the principal leptanilline and doryline section genera. In the diagnosis below, apparent apomorphic conditions are marked with an asterisk. Some of these are unique, others have appeared convergently in other ant groups, as discussed below. A comparison with Adetomyrma HNS appears after the diagnosis.
Tribe Amblyoponini HNS Forel, 1893
1. * Worker, queen. Anterior clypeal margin with a row of specialized, stout setae which give it a denticulate appearance (Figs 8 - 11, 14, 15); this may be accentuated by the setae arising from cuticular projections ( Mystrium HNS , some Amblyopone HNS ). The setae are secondarily reduced in some species, lost in Amblyopone mutica (Santschi) HNS .
2. *? Worker, queen. Mandibles overlapping at their tips when closed and typically, but not always, without distinct
3. * Worker, queen. Palp formula 5,3, with reductions therefrom (Brown, 1960).
4. Worker, queen. Twelve antennal segments, with uncommon secondary reduction to as few as 9 and 7.
5. Worker, queen. Antennal sclerite (torulus) raised medially and fusing with the overlapping frontal lobes.
6. * Worker. Compound eyes reduced or, less commonly, absent.
7. * Worker, queen. Eyes, when present, situated behind the middle of the sides of the head.
8. Worker. Promesonotal suture unfused.
9. * Worker. Mesonotum short and transverse, on a flattened mesosoma, shorter than the basal (= dorsal) face of the propodeum, usually much so.
10. * Worker, queen, male. Metacoxal cavities encircled by cuticle, the endpoints meeting broadly but not fused (Figs 19, 20). Fusion occurs as a secondary development within the tribe, in Onychomyrmex HNS and possibly others.
11. * Worker, queen. Metapleural gland orifice directed predominantly dorsally and posteriorly, on a curved surface mesad of a posterolateral swelling or plate; orifice plainly visible, as a thin crescentic cavity, in posterior view; no guard hairs arising from the posterolateral swelling (Figs 25, 26, 31, 32). The metapleural gland orifice of Myopopone HNS is somewhat divergent, opening laterally as well as posterodorsally.
12. * Worker, queen. Petiole (abdominal segment 2) shape characteristic: essentially apedunculate, with a steep, broad anterior face and a flat dorsal face that extends to the posterodorsal margin without descending into a distinct posterior face.
13. * Worker, queen, male. Abdominal sternum 2 with distinctive configuration: fused anteriorly with the tergum, but not fully fused posteriorly, i. e. with some free play between the sternite and adjacent tergite; posterior portion of sternum 2, i. e. section posterior to the region of definitive fusion, typically triangular or pyriform in shape (ventral view), its anterior apex occurring at a ' pinch point' marked by the approximate convergence of three structures: the laterotergite (see 14 below), the anterior termination of the (usually well developed and antero-dorsally undercut) ventral petiolar tooth, and the lower margins of the collar-like peduncular flange that encircles the tergum anteriorly (Fig. 37); very rarely (e. g. A. mutica HNS ) the petiolar tooth and peduncular flange absent.
14. *? Worker, queen, male. Abdominal tergum 2 with a distinct laterotergite which parallels the posterior portion of the sternum, forming a hinge joint with it; laterotergite broad posteriorly, narrowing anteriorly and terminating at the ' pinch point' near the lower margins of the collar-like peduncular flange (Fig. 37). Laterotergite reduced / lost in Prionopelta HNS (Fig. 38) and in most Onychomyrmex HNS .
15. *? Worker, queen, male. Petiole broadly attached to upper margins of abdominal segment 3 and helcial sternite broad. A significant narrowing of the broad dorsal attachment occurs only in the aberrant Amblyopone mutica HNS .
16. Worker, queen, male. Abdominal segment 3: ter-gosternal fusion of presclerites and postsclerites,
17. Worker, queen. Abdominal segment 4: differen-tiation of presclerites and tergosternal fusion of the entire segment.
18. * 7 Worker, queen, male. Absence of stridulatory file on abdominal tergum 4.
19. Worker. queen. Pygidium simple.
20. Worker, queen. Sting apparatus well developed, furcula present.
21. Worker, queen, male. Tibial spurs 1.2.2. with reductions therefrom.
22. * Worker, queen. Posterior metatibial spur stout subtriangular and curved (Hashimoto, 1991 b), its lower margin pectinate and upper margin barbulate.
23. Worker, queen, male. Tarsal claws simple.
Some of the starred features listed above (e. g. reduction and position of ine compound eyes: simple tarsal claws) appear repeatedly in other ant groups and hence have limited phylogenetic value. Other features, such as the absence of a stridulatory file and the broad attachment of the petiole to the succeeding segment, are possibly pleomorphic. This is almost certainly true of inc presence of a laterotergite on abdominal tergum 2 (also seen, for example, in Myrmecina HNS and some cerapachyines and other ponerines) but the particular ' pinch point' arrangement of
the laterotergite, the ventral petiolar tooth, and the peduncular * flange is more or less confined to Amblyoponini HNS . A ventral petiolar tooth and a peduncular flange are also seen in many other ponerines, but their shape and position in Amblyoponini HNS are distinctive. The petiolar tooth, in particular usually protrudes anteroventrally with the result that, in lateral profile, a notch-like incision is observed at its anterior junction with the sternum proper. Also compelling as synapomorphies are the specialized dentiform clypeal setae which appear to be unique to the tribe: the particular configuration of the metapleural gland which is not duplicated elsewhere: the shape of the petiole, which is rarely seen in other ants ( Typhlomyrmex rogenhoferi Mayr HNS and a few cerapachyines approach this condition); and the broad but primitively unfused connection between the sclerites encircling the metacoxal cavities. As far as known, all amblyoponines are cryptic in their foraging habits and specialized as predators on arthropods, especially centipedes, living in soil or rolling wood (Brown, 1960; Gotwald & Levieux, 1972; Hoelldobler & Wilson, 1986; Masuko, 1986; Ito, 1993) The shape of the mandibles, the powerful sting, and the dentiform clypeal (sometimes also labral) setae no doubt assist physically in prey capture (Brown, 1960), although the clypeal setae probably also have a sensory function.
Excluded from Amblyoponini HNS
1. Apomyrma HNS , known from the single species Apomyrma stygia Brown HNS , Gotwald & Levieux (1971), was originally placed in the Amblyoponini HNS but later transferred to the Leptanillinae HNS by Bolton (1990 a) and then given subfamily status, as Apomyrminae HNS in Baroni Urbani et al. (1992). The clypeal margin of the Apomyrma HNS worker is not adorned with specialised setae but there is a double row of similar, presumably analogous (see also Gotwald & Levieux, 1972) peg-like setae on the labrum. Note that these are different in shape than those of die Amblyoponini HNS (compare Figs. 8 - 11, 14 - 17). Nevertheless dentiform setae are round on both the labrum and the clypeal margin in Onychomyrmex HNS (Fig. 9) and some Amblyopone HNS , suggesting a possible link between the characters. Protanilla HNS , a leptanilline genus, also possesses a pair of stout setae on the labrum (Bolton, 1990 a) but none on the clypeus. Apomyrma HNS workers have no eyes and those of (he queen are placed in a posterior position on the head in agreement with Amblyoponini HNS . With respect to the remaining putative synapomorphies of Amblyoponini HNS , however, Apomyrma HNS exhibits differences; it has a fully closed metacoxal cavity surrounded by a dis-tinct annulus; metapleural gland orifice opening posterolaterally, under a dorsal cuticular flap; non-amblyoponine petiole shape: sharp constriction between the petiole and abdominal segment 3: and an isolated posterior petiolar sternite that is much reduced in size, although flanked by apparent laterotergites (Fig. 39). It must be admitted, however, that most of these conditions could be derived from those seen in Amblyoponini HNS ( Amblyopone mutica HNS provides an uncanny example of a petiole tending towards the Apomyrma HNS condition.) The same could be said for the horizontal toruli, forward-placed spiracle on abdominal tergum 3, palp formula of 2,2, and transverse sulcus behind the helcial sternite (see Bolton, 1990 a: 280). Hence a close relationship between these two cannot be ruled out. TheAmblyoponini HNS might even be paraphyletic relative to Apomyrma HNS (and Leptanillinae HNS ), although the morphology of the metapleural gland and gaster suggests otherwise.
2. The ponerine genus Typhlomyrmex HNS , placed in its own tribe but sometimes considered a possible relative of Amblyoponini HNS (Brown, 1974 b), has no specialized clypeal setae, a metapleural gland whose orifice opens dorsolateraly and is not visible in a strict posterior view (Figs 27, 33), open metacoxal cavities whose encircling cuticle does not overlap broadly (Fig. 21), and a differently shaped petiolar sternite (Fig. 40). Typhlomyrmex HNS also lacks a petiolar laterotergite and the broad attachment of the petiole to the postpetiole. Brown (1965) cites additional differences between Typhlomyrmex HNS and Amblyoponini HNS in wing venation and larval mandibles.