Corrieopone, Esteves & Fisher, 2021

Esteves, Flavia A. & Fisher, Brian L., 2021, Corrieopone nouragues gen. nov., sp. nov., a new Ponerinae from French Guiana (Hymenoptera, Formicidae), ZooKeys 1074, pp. 83-173 : 83

publication ID	https://dx.doi.org/10.3897/zookeys.1074.75551
publication LSID	lsid:zoobank.org:pub:0EE73BBC-E98A-46AC-B6B9-83BF15868F8B
persistent identifier	https://treatment.plazi.org/id/2B32773A-92E1-41B2-BC74-EC0B8ECD4A0A
taxon LSID	lsid:zoobank.org:act:2B32773A-92E1-41B2-BC74-EC0B8ECD4A0A
treatment provided by	ZooKeys by Pensoft
scientific name	Corrieopone
status	gen. nov.

Treatment

Corrieopone gen. nov.

Figs 7 View Figure 7 , 8 View Figure 8 , 37 View Figure 37 , 38 View Figure 38 , 39 View Figure 39 , 40 View Figure 40 , 41 View Figure 41 , 42 View Figure 42 , 43 View Figure 43 , 44 View Figure 44 , 52A View Figure 52 , 53 View Figure 53 , 54 View Figure 54 , 55 View Figure 55 , 56 View Figure 56 , 57 View Figure 57

Type species.

Corrieopone nouragues sp. nov., by present designation.

Diagnosis based on workers.

Medium-sized, slender Neotropical ants (TL 6.5-7.1 mm; Fig. 37 View Figure 37 ) with characters of Ponerinae (as in Fisher and Bolton 2016) and Ponerini (as in Schmidt and Shattuck 2014), in addition to the following (asterisks indicate putative autapomorphies):

1. Mandibles triangular, with distinct masticatory and basal margins; inserted at the anterolateral corners of the head (Fig. 38A, E View Figure 38 ).

2. Mandibles edentate (Fig. 38A, E View Figure 38 ).

3. Mandible devoid of any pit or sulcus: basolateral and dorsal pits and dorsolateral and dorsomasticatory sulci absent (Fig. 38A-C, E View Figure 38 ).

4. * Clypeus complex: In dorsal view, clypeus projected anteromedially as a broad, truncated prominence, overhanging the basal margins of the mandibles, and overlapping the basal portion of the masticatory margins of fully closed mandibles; anterior margin of the clypeal projection approximately as wide as the distance between the lateral arches of the toruli, devoid of stout setae or additional protrusions (Figs 37B View Figure 37 , 38A, E View Figure 38 ). In profile, clypeal projection with a broad anteroventral face (avf, Fig. 39A-D View Figure 39 ), which extends ventroposteriorly (i.e., obliquely) from the clypeal dorsal face in almost 90 degrees; the ventralmost point of this anteroventral face meets the “true” clypeal ventral face (= surface of the clypeal infold in Boudinot et al. 2021; vf, Fig. 39A-D View Figure 39 ). In anteroventral view, clypeal anteroventral face subrectangular (avf, Fig. 39D View Figure 39 ). Median area of the clypeus bulging (Fig. 37A, E View Figure 37 ); seen in profile, it ascends steeply from the clypeal anterior margin to the torular lobes, with posterior portion slightly convex (Fig. 38B, D View Figure 38 ).

5. In dorsal view, torular lobes closely approximated.

6. In dorsal view, torular lobes medium- to small-sized: not concealing the lateral arches of the toruli (Fig. 38F View Figure 38 ). Median and lateral arches of the torulus with discontinuous posterior margins (Fig. 38D View Figure 38 ).

7. In profile, torular lobes located at the dorsalmost part of a prominence formed by the clypeal median area and the frontal carinae (Fig. 38B View Figure 38 ).

8. Compound eyes small and located immediately anterior to the midline of the head (Fig. 38A, B, D View Figure 38 ).

9. Ocelli absent (Fig. 38A View Figure 38 ).

10. Labrum apically bilobed; with a long, acute cleft at the midpoint of its apical margin. Lobes broadly rounded apicolaterally and unarmed (Fig. 39E View Figure 39 ).

11. Palpal formula: 4,4 (four maxillary, four labial palpomeres; Fig. 39F View Figure 39 ).

12. Mesonotum rounded, dome-shaped in profile (Fig. 40A View Figure 40 ), rounded in dorsal view (Fig. 37A View Figure 37 ).

13. Notopleural suture distinct (Fig. 40A, B View Figure 40 ).

14. Metanotal sulcus deeply impressed (Fig. 40A View Figure 40 ).

15. Mesopleuron in profile divided into anepisternum and katepisternum (Fig. 40A, B View Figure 40 ).

16. Metathoracic spiracle concealed by a spiracular lobe (Fig. 40B View Figure 40 ).

17. Orifice of the metapleural gland round, opening posterolaterally on the metapleuron, with its ventral margin atop the posteriormost portion of the metapleural carina (Fig. 40C, D View Figure 40 ).

18. Metapleural longitudinal flange absent (Fig. 40C, D View Figure 40 ).

19. Propodeal dorsum devoid of a median longitudinal groove or impression.

20. Propodeum unarmed: without dorsoposterior projections (Fig. 40A View Figure 40 ).

21. In profile, propodeal lobe round, not surpassing posteriorly the dorsoposterior-most point of the rim of the propodeal foramen (Fig. 40C View Figure 40 ).

22. Propodeal spiracle slit-shaped (Fig. 40C View Figure 40 ).

23. Mesosternal process bidentate; metasternal process bilobate, long (Fig. 40F View Figure 40 ).

24. Metacoxal cavities open; cavities tightly encircled by cuticle, but cuticular annulus not fused (Fig. 40E View Figure 40 ; see comment in the following section).

25. Calcar of strigil with a basoventral lamella (Fig. 41A, B View Figure 41 ).

26. Probasitarsus with anterior and ventral faces densely vested with spatulate-costate setae (Fig. 41A View Figure 41 ), except for the shorter, spatulate-bicuspid setae present on the area immediately anterior to the comb of strigil (Fig. 41B View Figure 41 ).

27. Row of stout, spine-like setae present on the posterior face of the probasitarsal notch, parallel to the comb of strigil (Fig. 41B View Figure 41 ).

28. Two well-developed mesotibial spurs: anterior spur simple, posterior spur serrate (Fig. 41C, D View Figure 41 ).

29. Apparent metatibial gland cuticular patch present on the apicoposterior face of the metatibia, next to the posterior metatibial spur (Fig. 41E View Figure 41 ).

30. Two well-developed metatibial spurs: anterior spur simple, posterior spur pectinate (Fig. 41F View Figure 41 ).

31. Stout, spine-like setae absent from the dorsal face of mid- and hindlegs (Fig. 42A, B View Figure 42 ).

32. Ventral faces of the second, third, and fourth tarsomeres of fore-, mid-, and hindlegs with a paired row of stout, spine-like setae skirting the midline of each segment (Fig. 42C-E View Figure 42 ).

33. Pro-, meso-, and metapretarsi with simple claws (Fig. 42C-F View Figure 42 ).

34. Arolia indistinct (Fig. 42F View Figure 42 ).

35. Petiole sessile, with high, unarmed, conic, scale-like node (i.e., tergite narrow in profile and dorsal view; Figs 37A, C View Figure 37 , 43A View Figure 43 ).

36. Petiolar tergite with anteroventral lateral carina (= lateral, dorsoventral carina in Ward 1990, character 29); posteroventral portion of tergite strigate (Fig. 43A View Figure 43 ).

37. Petiolar laterotergite distinct (Fig. 43A View Figure 43 ).

38. Proprioceptor zone on anterior disc of petiolar sternite shaped as a large circular area (Fig. 43B View Figure 43 ).

39. Petiolar sternite without posterior spatulate projection (Fig. 43B View Figure 43 ); articulation with helcium visible in ventral view.

40. Helcium infra-axial: positioned ventrad the midheight of the anterior face of abdominal segment III (Fig. 43C View Figure 43 ).

41. Prora present as a lip-shaped, transverse projection on the anterior portion of the abdominal poststernite III (Fig. 43C, D View Figure 43 ); projection not extending anteriorly to the area between the ventroposterior margins of helcial tergite (Fig. 43D View Figure 43 ).

42. Abdominal segment IV tubular: tergite and sternite with similar lengths; tergite not arched (Fig. 43C View Figure 43 ).

43. Presclerites of abdominal segment IV forming an even surface with postsclerites: girdling constriction absent (Fig. 43C View Figure 43 ).

44. Stridulitrum present on abdominal pretergite IV, small (Fig. 43E, F View Figure 43 ).

45. Pygidium devoid of stout, spine-like setae or spine-like microtrichia; dorsal face convex.

46. * Ventral face of hypopygium longitudinally concave (Fig. 44A, B View Figure 44 ). Longitudinal carina present at midline of concavity’s posterior portion, skirted laterally by stout, hook-shaped setae (Fig. 44C, D View Figure 44 ).

47. Hypopygium posterolateral region without stout, spine-like setae or spine-like microtrichia.

Comments on worker characters

The enumeration below corresponds to character numbers presented above.

1. The mandibles articulate with the anterolateral corners of the head in virtually all Ponerinae and are either triangular or subtriangular in most genera (N = 29). Other mandibular shapes with little intrageneric variation are the oblique ( Boloponera , Buniapone , Dinoponera , Iroponera , Plectroctena , and Promyopias ), pitchfork-like ( Belonopelta , Emeryopone , Thaumatomyrmex ), elongate sub-oblique ( Streblognathus ), and scythe-shaped ( Harpegnathos ). The shape of the mandibles varies from triangular to elongate-triangular in Centromyrmex ; from triangular to subtriangular, to oblique in Cryptopone ; from subtriangular to oblique, to falcate, to bizarre forms in-between in Leptogenys and Myopias ; from sub-oblique to falcate in Psalidomyrmex ; and from subtriangular to oblique in Simopelta . Anochetus and Odontomachus are the only ponerines in which the mandibles insert near the midline of the anterior margin of the head.

2. The mandible of Corrieopone is completely edentate (i.e., devoid of any teeth, denticle, or projected apex). To our knowledge, this condition is virtually absent in other Ponerinae , apart from some species of Leptogenys (see Arimoto and Yamane 2018) and Platythyrea (see Fisher and Bolton 2016).

3. Ponerine may present mandibles ornamented with pits and sulci, which are relatively good diagnostic characters to genera.

3.1. The basolateral pit (= basal pit in Fisher and Bolton 2016) is a round to oblong impression on the basal portion of the lateral face of the mandible. It occurs in most Brachyponera species (excluding species from Borneo, Bali, Krakatau, and Sumatra, according to Yamane 2007), Cryptopone [except for C. guianensis and, based on its original description, C. mirabilis (Mackay & Mackay), as far as we know], and Euponera (see Schmidt and Shattuck 2014). The pit is also present in Fisheropone ambigua (Fig. 45A, B View Figure 45 ), which disagrees with Schmidt and Shattuck (2014) and Fisher and Bolton (2016).

3.2. The dorsal pit resembles the basolateral pit, although more elongated and impressed on the dorsal face of the mandible. In Ponerinae , it is only present in Dolioponera , Euponera fossigera Mayr (see Mayr 1901), Hagensia , and Iroponera (Fig. 45C View Figure 45 ; contrary to Schmidt and Shattuck 2014).

3.3. The dorsolateral sulcus runs obliquely along the mandible, from the basal portion of the dorsal face towards the lateral face. It is widespread among the Ponerinae and may be shallowly or deeply impressed, restricted to the basal portion of the mandible, or present along almost the entire lateral face of the mandible. The sulcus is consistently present in species of Asphinctopone , Boloponera , Buniapone , Centromyrmex , most (perhaps all) Ectomomyrmex , Feroponera , Loboponera , Myopias , Odontoponera , Paltothyreus , Phrynoponera , Plectroctena , Promyopias , Psalidomyrmex , Pseudoponera , and Streblognathus (see Suppl. material 3: Table S3; Schmidt and Shattuck 2014; Fisher and Bolton 2016). On the other hand, the presence of this character varies among species of other genera, such as Bothroponera (present in B. crassa , absent in B. pachyderma ), Dinoponera (weakly present in D. lucida ; absent in D. longipes ), Leptogenys (see Bolton 1975), Mesoponera (only present in M. subiridescens ), Neoponera [present in N. fauveli (Emery), weakly impressed in N. apicalis , absent in N. fisheri ], Platythyrea (present in P. punctata , absent in P. turneri ), and Pseudoneoponera (present in P. porcata , absent in P. denticulata ). Contrary to Schmidt and Shattuck (2014), the sulcus is distinct in all Rasopone species examined here (Fig. 45D View Figure 45 ); although short and constrained to the basal region of the mandibles, it is also present in Austroponera castanea (Fig. 45E View Figure 45 ). Among the Pachycondyla species examined, the sulcus is a shallow and short basal impression save in P. lenis , where it is absent. In Euponera sikorae (and in all other Malagasy species, according to Rakotonirina and Fisher 2013), the lateral face of the mandible bears a longitudinal sulcus that runs apicad from the lateral margin of the dorsal mandibular articulation. Whether this sulcus is present in Euponera species occurring in other bioregions remains unknown, but it is absent in at least two Afrotropical species, E. brunoi and E. sjostedti .

3.4. Plectroctena species present a sulcus that skirts the mandibular masticatory margin dorsally ( Fisher and Bolton 2016), which we refer to as the dorsomasticatory sulcus.

Among the specimens examined, the following taxa present mandibles devoid of any pit or sulcus, like Corrieopone : Anochetus angolensis , A. emarginatus , Belonopelta deletrix , Bothroponera cariosa , B. pachyderma , B. talpa , Cryptopone guianensis , Diacamma ceylonense , Dinoponera longipes , Emeryopone buttelreepeni , Harpegnathos saltator , Hypoponera punctatissima , Mayaponera , Megaponera analis , Mesoponera ambigua , M. australis , M. caffraria , M. elisae rotundata , M. melanaria macra , M. papuana , M. rubra , Neoponera commutata , N. fisheri , N. laevigata , N. luteola , N. verenae , N. villosa , Odontomachus bauri , Ophthalmopone berthoudi , Pachycondyla lenis , Parvaponera darwinii madecassa , Platythyrea turneri , Ponera alpha , P. pennsylvanica , Simopelta oculata , S. transversa , Thaumatomyrmex fraxini , and T. zeteki .

6. We assessed the size of the torular lobes across taxa examined according to the degree of connection between median and lateral arches of torulus (as in Keller 2011, character 08). Among material examined, the lobes are anteriorly and posteriorly continuous with the lateral torular arches in taxa whose antennal sockets are largely exposed in full-face view ( Belonopelta , Leptogenys , and Ophthalmopone ). On the other extreme, hypertrophied lobes are anteriorly and posteriorly discontinuous with the lateral arches of the torulus (as in Boloponera , Bothroponera sensu stricto group, Loboponera , Platythyrea punctata , Plectroctena , and Psalidomyrmex ). Like Corrieopone , most ponerines present intermediate-sized torular lobes that are anteriorly continuous and posteriorly discontinuous with respective lateral arches. In this latter state, the lobes may conceal entirely or partially the lateral arches of the torulus.

For the record, the torular lobes in Bothroponera sulcata species-group members (sensu Schmidt and Shattuck 2014) resemble those of Corrieopone , except for B. henryi (Donisthorpe) (see specimen CASENT0902482) and B. zumpti Santschi (see CASENT0922370), where the lobes are hypertrophied as in the Bothroponera sensu stricto group.

9. Ocelli are invariably present on Harpegnathos workers; it is absent in the worker caste of other Ponerinae , apart from occasional workers.

11. A count of four maxillary and four labial palpomeres is consistently present in Buniapone , Dinoponera , Hagensia , Harpegnathos , Mayaponera (note that we did not examine M. longidentata ), Megaponera , Odontoponera , Ophthalmopone , Paltothyreus , Phrynoponera , Promyopias , Streblognathus (see Suppl. material 3: Table S3; Bolton 2003; Fisher and Bolton 2016). We are hesitant to affirm that this is also the case in Neoponera , Pachycondyla , Pseudoneoponera , and Rasopone . These characters usually have been neglected in taxonomic reviews and descriptions of new species, and we did not examine every species in these genera. Yet, in every species we did examine ( Neoponera aenescens , N. apicalis , N. bugabensis , N. carinulata , N. commutata , N. crenata , N. curiosa , N. dismarginata , N. fisheri , N. foetida , N. globularia , N. insignis , N. inversa , N. laevigata , N. luteola , N. moesta , N. obscuricornis , N. schoedli , N. striadinodis , N. unidentata , N. verenae , N. villosa , Pachycondyla crassinoda , P. harpax , P. impressa , P. lattkei , P. lenis , P. procidua , P. striata , Pseudoneoponera porcata , P. tridentata , Rasopone costaricensis , R. cryptergate s, R. cubitalis , R. guatemalensis , R. panamensis , R. pluviselva , and R. Rasopone politognatha ), the palpal formula was 4,4. The count is also 4,4 in some species of genera with variable palpal formulae, such as Anochetus , Bothroponera , Leptogenys , Mesoponera , and Myopias (see Suppl. material 3: Table S3; Willey and Brown 1983; Bolton 2003; Fisher and Bolton 2016).

While conducting this study, we noticed that the palpal formula of some taxa was incorrectly reported or missing in the pertinent literature. Thus we correct or update the record here. Contrary to Fisher and Bolton (2016), the palpal formula in Dolioponera fustigera is 3,3 (not 2,2, as previously stated; Fig. 45F View Figure 45 ), and 3,3 in Fisheropone ambigua (not 3,2; Fig. 45G, H View Figure 45 ). Contrary to Bolton (2003), Keller (2011), and Fisher and Bolton (2016), the palpal formula in Loboponera obeliscata is 2,3 (not 2,2; Fig. 45I, J View Figure 45 ); the formula in L. vigilans was correctly reported as 2,2 by these authors (verified on specimen CASENT0003102). We also report for the first time the palpal formula in Boloponera ikemkha (2,3; Fig. 45K, L View Figure 45 ), Cryptopone hartwigi (3,3; Fig. 46A, B View Figure 46 ), and Simopelta transversa (2,2; Fig. 46C View Figure 46 ). The specimen of Thaumatomyrmex atrox examined by Keller (2011) was later determined to be T. fraxini by D’Esquivel et al. (2017) - note that the specimen is not in the list of material examined by the latter authors, but some SEM images taken by Keller (2011; specimen ANTWEB1008597) were used to illustrate the new taxon description. One detail not mentioned by D’Esquivel et al. (2017) was the palpal formula of T. fraxini , which is 3,3 (Fig. 46D View Figure 46 ; previously reported for T. atrox by Keller 2011). Finally, although Fisher and Bolton (2016) stated that the palpal formula in Parvaponera darwinii madecassa is unknown, Brown (1963) provided the correct count, which is 4,3 (Fig. 46E View Figure 46 ).

12-14. In Corrieopone , the mesonotum is dome-shaped in profile, with a round dorsal margin that is discontinuous with the outline of the pronotum (i.e., it is slightly higher than the pronotum). The promesonotum is much higher than the propodeum, and a deeply impressed metanotal sulcus separates the two. A distinct notopleural suture delimits the mesonotum from the mesopleuron. In dorsal view, the mesonotum is round. Taxa that bear some resemblance to Corrieopone in this combination of characters are: several Anochetus species (e.g., A. altisquamis Mayr, specimen CASENT0915154; A. armstrongi McAreavey, CASENT0902449; A. brevis Brown, CASENT0902439), Asphinctopone , Austroponera [except A. rufonigra (Clark), CASENT0249178], Brachyponera , Euponera sikorae , Fisheropone ambigua , Hagensia , some Hypoponera [e.g., H. foreli (Mayr), CASENT0173714; H. herbertonensis (Forel), CASENT0907320; H. mesoponeroides (Radchenko), CASENT0917250], some Leptogenys (e.g., L. borivava Rakotonirina & Fisher, CASENT0430091; L. ixta , L. peruana , L. sonora ), Mayaponera , Megaponera analis (dome-shaped mesonotum in larger specimens, as CASENT0781129), most Mesoponera (e.g., M. ambigua , M. australis , M. caffraria , M. elisae rotundata , M. melanaria , M. papuana , M. rubra , M. subiridescens ), several Myopias [e.g., M. castaneicola (Donisthorpe), CASENT0902520; M. chapmani Willey & Brown, CASENT0902533; M. latinoda (Emery), CASENT0270592], several Neoponera (e.g., N. apicalis , N. aenescens , N. fisheri , N. schoedli , N. villosa ), some Odontomachus (e.g., O. bauri ; O. laticeps Roger, CASENT0904008; O. spissus Kempf, CASENT0281868), Odontoponera transversa , Ophthalmopone , Rasopone rupinicola , R. cubitalis , and Streblognathus .

16. The spiracular lobe is present in most ponerine genera; it is present in Boloponera ikemkha (CASENT0254321) and B. vicans (CASENT0401737), contrary to Fisher and Bolton (2016).

The lobe is absent in Dolioponera , Fisheropone , some Loboponera species, the Afrotropical and Malagasy Hypoponera , H. punctatissima , Simopelta oculata , S. transversa , and Thaumatomyrmex fraxini (Suppl. material 3: Table S3; Fisher and Bolton 2016). Interestingly, while the lobe is absent in the Afrotropical Cryptopone (according to Fisher and Bolton 2016; here confirmed on C. hartwigi ), it is present in the Neotropical C. gilva (ANTWEB1008514), C. guianensis (ANTWEB1008565), C. holmgreni (ECOFOG-IT14-0276-07), and C. pauli (CASENT0637806).

18. The metapleural longitudinal flange is a carina that extends along the metapleuron in profile, with its posterior end immediately dorsad the metapleural gland orifice. When well-developed, it projects laterad or ventrolaterad and may overhang the gland orifice (as defined by Keller 2011: character 62). In Simopelta species (ANTWEB1008589), this flange is strongly projected ventrolaterad and overlaps the gland orifice.

19. The propodeal dorsum presents a well-delimited, narrow, median longitudinal groove in Psalidomyrmex ( Fisher and Bolton 2016; ANTWEB1008585). In addition, a vestigial longitudinal groove may be present on the propodeal dorsal face of specimens of the Plectroctena minor group (see Bolton and Brown 2002). The propodeal dorsum is transversely concave along its entire length, or only posteriorly, in Hagensia , Mayaponera (CASENT0249137), several Mesoponera (CASENT0249194), and Pseudoponera (CASENT0923115).

20. Most ponerine genera present an unarmed propodeum. In profile, the propodeal dorsoposterior corner bears acute projections in several Anochetus species (e.g., CASENT0902431, CASENT0815182, CASENT0746783), Phrynoponera (CASENT0178230), Streblognathus (ANTWEB1008591), some Platythyrea (CASENT0281867, CASENT0900569, CASENT0903799), and Pseudoneoponera bispinosa (Smith). Also, some species of the Loboponera vigilans group (CASENT0003111), Plectroctena Plectroctena minor group (CASENT0915285), and P. mandibularis group (CASENT0102947) present the lateral margin of the propodeal declivity with a lamella that is dorsally toothed; in other species of the L. vigilans group, the lateral margin of the propodeal declivity is toothed dorsally, but the lamella is absent (CASENT0003098; see Bolton and Brown 2002).

22. In general, the shape of the propodeal spiracle is constant within genus in Ponerinae . A slit-shaped spiracle (i.e., external atrial opening> 2 × longer than wide; as in Keller 2011, character 65) is present in Asphinctopone , Austroponera , Bothroponera , Buniapone , Corrieopone , Diacamma , Dinoponera , Ectomomyrmex , Euponera , Feroponera , Fisheropone , Hagensia , Harpegnathos , Megaponera , Odontoponera , Ophthalmopone , Pachycondyla , Paltothyreus , Phrynoponera , Pseudoneoponera , and Streblognathus . A round to oval spiracle occurs in Belonopelta , Boloponera , Dolioponera , Emeryopone , Hypoponera , Iroponera , Loboponera , Mayaponera , Myopias , Odontomachus , Plectroctena , Ponera , Promyopias , Psalidomyrmex , Rasopone , Simopelta , and Thaumatomyrmex . The propodeal spiracle is round to oval in most Brachyponera species but varies from oval to slit-shaped in B. atrata and B. sennaarensis .

24. We classified the metacoxal cavities as closed or open by integrating the definition given by Keller (2011: character 69) with that in Fisher and Bolton (2016; see also Bolton 2003: character 41). If closed, the medial surface of the metacoxal acetabulum does not have a fenestra, and thus, the metacoxal cavity is not connected internally with the propodeal foramen; the annulus externally encircling the cavity is fused. If open, the internal medial surface of the cavity is fenestrate and connects with the propodeal foramen, and the annulus is unfused. In this case, the annulus may encircle the cavity with its free ends overlapping next to the propodeal foramen; or there may be a gap in the annulus.

An unfused cuticular annulus tightly encircles the metacoxal cavities in most Ponerinae we dissected. This condition occurs in Phrynoponera pulchella specimens from Kenya (CASENT0178203, CASENT0178204, CASENT0217125), which is in accord with Bolton and Fisher (2008a). However, a specimen from Tanzania possesses an annular gap in both metacoxal cavities (Fig. 46H View Figure 46 ); we are uncertain if those were natural or dissection artifacts, as unfortunately, only one specimen from that population was available to us. Metacoxal cavities also present an annular gap in Phrynoponera transversa and Platythyrea punctata . On the other hand, the annulus is fused and uninterrupted in Harpegnathos saltator , Platythyrea cribrinodis (Fig. 46F View Figure 46 ; contrary to Fisher and Bolton 2016), and Myopias darioi (Fig. 46G View Figure 46 ).

25. The calcar of strigil presents a basoventral lamella in most Ponerinae evaluated (as in Keller 2011, character 74). For clarification, we consider the calcar to present a small lamella in Loboponera obeliscata (Fig. 46I View Figure 46 ) and that it is entirely pectinate in Platythyrea punctata and P. turneri (specimens ANTWEB1008574 and ANTWEB1008575, respectively), which is contrary to Keller (2011). The lamella is also absent in Boloponera , Brachyponera sennaarensis , Diacamma ceylonense , Dolioponera fustigera , Emeryopone buttelreepeni , Harpegnathos saltator , Hypoponera punctatissima , Leptogenys ixta , L. peruana , L. pucuna , L. sonora , L. wheeleri , Loboponera vigilans , Mesoponera ambigua , M. elisae rotundata , Myopias darioi , Platythyrea cribrinodis , Ponera alpha , P. pennsylvanica , Promyopias silvestrii , Simopelta oculata , S. transversa , Thaumatomyrmex fraxini , and T. zeteki .

27. A row of stout, spine-like setae occurs along the posterior face of probasitarsal notch, parallel to the comb of strigil, in most Ponerinae taxa examined. We adopted the definition of “row” analogous to that of a line, whose existence requires at least two points in space. Thus, the row is present if two or more spine-like setae are aligned longitudinally along the posterior face of the probasitarsal notch; less than two setae make the row absent. Among material examined, the row is absent on the posterior face of the probasitarsal notch in Dolioponera fustigera , Leptogenys , Myopias darioi , and Thaumatomyrmex fraxini .

28. Most Ponerinae taxa present two mesotibial spurs. Among taxa examined, the anterior spur is simple, and the posterior spur is serrate in Brachyponera chinensis , B. croceicornis , B. lutea , B. luteipes , B. obscurans , Dinoponera longipes , D. lucida , Hagensia havilandi marleyi , Harpegnathos saltator , Leptogenys peruana , L. pucuna , and Ophthalmopone berthoudi .

For the record, only one mesotibial spur is visible under a stereomicroscope in Asphinctopone silvestrii and Fisheropone ambigua ; however, SEM images reveal that a vestigial anterior spur is present in both taxa (Fig. 47C, E View Figure 47 ); A. differens also seems to bear a minute anterior spur on the mesotibia.

29. The metatibial gland has been confirmed in Bothroponera tesseronoda (Emery), several species of Diacamma Mayr, Harpegnathos saltator , Neoponera crenata , N. marginata (Roger), Paltothyreus tarsatus , Pseudoneoponera rufipes (Jerdon), and P. tridentata (see Hölldobler et al. 1996; Billen 2009). The gland is associated with a cuticular pore plate, which may be covered by a brush of stouter, distinctly-shaped setae (as in Diacamma and Paltothyreus tarsatus ), or it may be an oblong, glabrous, distinctly colored, smooth, or distinctly sculptured cuticular patch ( Hölldobler et al. 1996). Here, we list taxa that present the latter condition, which is what is seen in Corrieopone . Among the taxa examined, the cuticular patch is present on the apicoposterior face of the metatibia in Bothroponera crassa , B. silvestrii , Brachyponera croceicornis , B. lutea , B. luteipes , B. obscurans , B. sennaarensis , Cryptopone hartwigi , Ectomomyrmex javanus , Emeryopone buttelreepeni , Euponera sikorae , E. sjostedti , Feroponera ferox , Hagensia havilandi marleyi , Mesoponera caffraria , Pseudoneoponera porcata , and P. tridentata .

30. Most Ponerinae taxa present two metatibial spurs. Here, all specimens examined present a pectinate posterior spur, and in the majority, the anterior spur is simple, as in Corrieopone (see Suppl. material 3: Table S3).

For the record, Schmidt and Shattuck (2014: 185) and Fernandes and Delabie (2019: 411) stated that Cryptopone species present one metatibial spur (excluding C. guianensis , and in the case of the latter authors, also C. pauli ). Those assertions are puzzling, as several Cryptopone species have two spurs on the metatibia [e.g., C. arabica Collingwood & Agosti, C. gilva , C. holmgreni (Wheeler), C. ochracea Mayr; see Fig. 47A View Figure 47 ; Kempf 1961b; Bernard 1967; Collingwood and Agosti 1996]. Belonopelta deletrix has two spurs on the metatibia (Fig. 47B View Figure 47 ), as correctly stated by Baroni Urbani (1975), but contrary to Schmidt and Shattuck (2014). Asphinctopone silvestrii presents a minute additional metatibial spur (Fig. 47D View Figure 47 ), anterior to the much larger pectinate spur; A. differens seems to present the same. The metatibia of Fisheropone ambigua , which bears a single spur, long and pectinate, also presents a fovea where the anterior spur would be located (Fig. 47F View Figure 47 ).

31. Taxa without stout, spine-like setae on the dorsal face of the mid- and hindlegs are Anochetus angolensis , A. emarginatus , Asphinctopone differens , As. silvestrii , Belonopelta deletrix , Boloponera ikemkha , B. vicans , Bothroponera silvestrii , Brachyponera chinensis , B. croceicornis , B. luteipes , B. obscurans , B. sennaarensis , Diacamma ceylonense , Dolioponera fustigera , Emeryopone buttelreepeni , Euponera sikorae , E. sjostedti , Hagensia havilandi marleyi , Harpegnathos saltator , Iroponera odax , Leptogenys ixta , L. peruana , L. pucuna , L. sonora , L. wheeleri , Loboponera obeliscata , L. vigilans , Mayaponera conicula , Myopias darioi , M. maligna , Neoponera bugabensis , N. carinulata , N. cavinodis , N. crenata , N. dismarginata , N. fiebrigi cf., N. fisheri , N. foetida , N. globularia , N. insignis , N. inversa , N. luteola , N. moesta , N. obscuricornis , N. striadinodis , N. unidentata , N. villosa , Odontomachus bauri , Odontoponera transversa , Ophthalmopone berthoudi , Platythyrea cribrinodis , P. punctata , P. turneri , Simopelta oculata , S. transversa , Thaumatomyrmex fraxini , and T. zeteki .

Contrary to Fisher and Bolton (2016), Fisheropone ambigua presents stout, spine-like setae on the dorsal face of the mesobasitarsus (Fig. 48A View Figure 48 ).

33. We categorized the shape of pro-, meso-, and metapretarsal claws according to the presence, location, and number of acute projections on their inner margins (modified from Keller 2011, character 82). A simple pretarsal claw is devoid of prominences (Fig. 48B View Figure 48 ) or presents a blunt basal angle or rounded swell (Fig. 48C View Figure 48 ), and is found in the following taxa: Anochetus angolensis , A. emarginatus , Asphinctopone differens , A. silvestrii , Austroponera castanea , Belonopelta deletrix , Boloponera ikemkha , B. vicans , Bothroponera crassa , B. silvestrii , Brachyponera chinensis , B. croceicornis , B. lutea , B. luteipes , B. obscurans , B. sennaarensis , Centromyrmex brachycola , C. decessor , C. ereptor , C. raptor , Cryptopone gilva , C. guianensis , C. hartwigi , Diacamma ceylonense , Dolioponera fustigera , Ectomomyrmex javanus , Emeryopone buttelreepeni , Euponera brunoi , E. sikorae , E. sjostedti , Feroponera ferox , Fisheropone ambigua , Hypoponera punctatissima , Iroponera odax , Loboponera obeliscata , L. vigilans , Mayaponera becculata , M. cernua , M. conicula , M. constricta , Mesoponera ambigua , M. australis , M. caffraria , M. elisae rotundata , M. melanaria macra , M. papuana , M. rubra , M. subiridescens , Myopias darioi , M. maligna , Neoponera aenescens , N. apicalis , N. bugabensis , N. cavinodis , N. crenata , N. dismarginata , N. eleonorae , N. fiebrigi cf., N. fisheri , N. foetida , N. globularia , N. insignis , N. inversa , N. laevigata , N. luteola , N. obscuricornis , N. schoedli , N. striadinodis , N. unidentata , N. verenae , N. villosa , Odontomachus bauri , Odontoponera transversa , Pachycondyla lattkei , Parvaponera darwinii madecassa , Plectroctena strigosa , Ponera alpha , P. pennsylvanica , Psalidomyrmex procerus , Pseudoneoponera porcata , Pseudoponera gilberti , P. stigma , Rasopone costaricensis , R. cryptergates , R. cubitalis , R. guatemalensis , R. panamensis , R. pluviselva , R. politognatha , Simopelta oculata , S. transversa , and Streblognathus peetersi .

A claw with a basal tooth bears an acute prominence on the basal third of its inner margin (Fig. 12B View Figure 12 ). The basal tooth overhangs the outline of the inner margin of the claw and may bear several small, acute projections (Fig. 48D View Figure 48 ). This type of claw occurs in Bothroponera cariosa , B. pachyderma , B. talpa , Mayaponera arhuaca , M. pergandei , Megaponera analis , Neoponera carinulata , N. commutata , N. moesta , Ophthalmopone berthoudi , Pachycondyla crassinoda , P. harpax , P. impressa , P. lenis , P. procidua , P. striata , Phrynoponera pulchella , P. transversa , and Pseudoneoponera tridentata . A claw with a preapical tooth has an acute projection rising from the apical two-thirds of its inner margin (Fig. 48E View Figure 48 ). It is present in Dinoponera longipes , D. lucida , Hagensia havilandi marleyi , Harpegnathos saltator , Paltothyreus tarsatus , Platythyrea cribrinodis , P. punctata , P. turneri , Thaumatomyrmex fraxini , and T. zeteki . Pectinate pretarsal claws are shaped like a comb and are unique to Leptogenys among Ponerinae (Fig. 48F View Figure 48 ).

Finally, we found shape variations among the claws of the fore-, mid-, and hindlegs only in Buniapone amblyops and Promyopias silvestrii . These taxa present a propretarsal claw with a long basal tooth, while their meso- and metapretarsi claws are simple (Fig. 48G-I View Figure 48 ). This condition contradicts Bolton and Fisher (2008b) and Fisher and Bolton (2016), who stated the claws are simple in P. silvestrii .

34. The arolium was absent or reduced to a membranous cuticular flap between the pretarsal claws of most taxa examined (Figs 12B View Figure 12 , 48D-I View Figure 48 ). Exceptions were Belonopelta deletrix , Diacamma ceylonense , Harpegnathos saltator , Iroponera odax , Mayaponera , Neoponera , Parvaponera darwinii madecassa , Platythyrea cribrinodis , P. punctata , P. turneri , Rasopone guatemalensis , Simopelta oculata , S. transversa (Fig. 48C View Figure 48 ), Thaumatomyrmex fraxini , and T. zeteki .

Contrary to Schmidt and Shattuck (2014), we consider that Mayaponera constricta has indistinct arolia like all its congeners (Fig. 48B View Figure 48 ). In this species, the arolium is reduced to a cuticular flap, although more developed than what we classified as indistinct in other taxa. However, this feature was not noticeable under our stereo microscope.

35. The petiolar node of Corrieopone lacks a spine-like or any other acute projection and is narrow in profile, with its anterior and posterior surfaces tapering to an insignificant dorsal surface.

Ponerine taxa with an unarmed scale-like petiole are: several Anochetus (see CASENT0915154); Asphinctopone (CASENT0915481); Austroponera (FOCOL0965); Brachyponera (CASENT0915660); Buniapone (CASENT0903944); some Cryptopone (ANTWEB1008000); some Ectomomyrmex (CASENT0907270); Euponera fossigera species group [viz.: E. brunoi , E. fossigera Mayr (SAM-HYM-C002649B), E. malayana (Wheeler), E. sharpi Forel, E. wroughtonii Forel, E. wroughtonii crudelis Forel, and probably also E. sakishimensis (Terayama)]; Fisheropone (CASENT0906215); Hagensia (CASENT0256487); several Hypoponera (CASENT0281911); some Leptogenys (CASENT0902609); Mayaponera (USNMENT00442104); Mesoponera (CASENT0249169); some Neoponera (ANTWEB1014009); very few Odontomachus (CASENT0281868); very few Pachycondyla (UFV-LABECOL-000002); some Parvaponera (CASENT0915276); some Ponera (CASENT0235336); and Pseudoponera [CASENT0902509; except P. pachynoda (Clark), ANTWEB1008183].

36. The strigation on the posteroventral portion of the petiolar tergite of Corrieopone resembles that of Asphinctopone (CASENT0178221).

39. The spatulate projection rises from the posterior portion of the petiolar sternite and extends posteriad, overlapping either partially or entirely the remaining sternite. This definition departs from Keller (2011, character 111) in not requiring (a) the projection to extend beyond the posterior margin of the petiolar sternite and (b) close proximity between projection and remaining sternite. The reason is the character varies continuously, which invalidates both requirements. The variation comprises projections that are long and conceal the helcial sternite almost completely to partially (as in Phrynoponera pulchella , specimen ANTWEB1008573, and Platythyrea punctata , respectively; Fig. 49A View Figure 49 ); those that reach or almost reach the posterior margin of the petiolar sternite (as in Platythyrea turneri , Fig. 49B View Figure 49 ); and those even shorter (as in Asphinctopone silvestrii ; Fig. 49C View Figure 49 ). In addition, projections may tightly envelop the remaining sternite (as in Phrynoponera pulchella and Platythyrea punctata , ANTWEB1008574; Fig. 49D View Figure 49 ); or a slight gap may be present [as in Platythyrea turneri , specimen ANTWEB1008575, and Phrynoponera gabonensis ( André); Fig. 49E View Figure 49 ]; or the gap may be slightly broader (as in Rasopone jtl030; Fig. 49F View Figure 49 ) to much broader (as in Streblognathus peetersi ; Fig. 49G View Figure 49 ).

Thus, according to our definition, the spatulate projection of the petiolar sternite is present in Austroponera , Asphinctopone , Brachyponera , Megaponera analis , Ophthalmopone , Phrynoponera , Platythyrea , Rasopone , and Streblognathus ; see also Fisher and Bolton (2016) for the description of the sternite in some of these taxa. The petiolar sternite in Belonopelta deletrix , immediately anterior to its posterior margin, is projected ventrad and slightly posteriad; the same seems to happen in Thaumatomyrmex fraxini . According to the species redescription given by Mackay and Mackay (2010), Neoponera magnifica may also present the character.

40. Like Corrieopone , most ponerine genera present an infra-axial helcium (i.e., positioned ventrad the midheight of the anterior face of abdominal segment III; see Keller 2011, character 114). A few taxa examined present an axial helcium (i.e., positioned at midheight of the anterior face of abdominal segment III): Boloponera , Buniapone , Centromyrmex , Cryptopone (except C. guianensis ), Dolioponera , Feroponera , Iroponera , Platythyrea , and Promyopias .

41. As far as we know, the prora is present and well-developed in most Ponerinae . It is usually indistinct in Platythyrea , but contrary to Fisher and Bolton (2016), it is well-developed in some species [as in P. lamellosa (Roger); Fig. 49H View Figure 49 ], and weakly projected but still visible in others [as in P. pilosula (Smith); Fig. 49I View Figure 49 ]. According to Schmidt and Shattuck (2014), the prora is absent in Iroponera , but SEM images of I. odax reveal that it is present, albeit weakly projected (Fig. 49J View Figure 49 ). According to Fisher and Bolton (2016), the trait is absent in Dolioponera , which agrees with what we saw in most specimens. However, SEM images of specimen ANTWEB1008521 show a weak projection on the anterior face of abdominal sternite III; we are unsure whether that detail is natural or an image artifact.

We consider the prora present in Mayaponera and Rasopone , in disagreement with Longino and Branstetter (2020). The trait, although small, is distinct in the profile view of M. becculata , M. conicula , M. constricta , and R. panamensis . In the first two species and R. panamensis , the prora rises from the anterior portion of the abdominal poststernite III (see specimens CASENT0249130, CASENT0644252). In M. constricta , it projects from the area in between the ventral margins of the helcial tergite arch and the anterior portion of the abdominal poststernite III (Fig. 49K View Figure 49 ) and resembles the condition seen in Dinoponera , Pachycondyla , and Streblognathus (ANTWEB1014000, CASENT0249148, and Fig. 49G View Figure 49 , respectively). In M. arhuaca , M. cernua , M. pergandei , and other Rasopone species, the prora is indistinct in the profile view of undissected specimens, for it is a minute prominence located in the area between the ventral margins of the helcial tergite (as in Brachyponera , Fig. 49L View Figure 49 ), and may be fused entirely with it (as described for Phrynoponera by Bolton and Fisher 2008a).

43. We considered the girdling constriction present if the surface between pre- and postsclerites of abdominal segment IV was interrupted by a shallow or deep impression on the integument. A line, if present, only constituted a constriction if the integument was depressed.

Among taxa examined, the constriction is absent in Asphinctopone , Brachyponera sennaarensis (weakly impressed in other species), Corrieopone , Mesoponera (except M. caffraria ), Odontomachus , Odontoponera , Simopelta , Streblognathus , and Thaumatomyrmex fraxini .

44. The stridulitrum is consistently present on abdominal pretergite IV in Austroponera , Belonopelta , Dinoponera , Harpegnathos , Megaponera , Neoponera , Odontoponera , Ophthalmopone , Streblognathus , and Thaumatomyrmex . The trait’s occurrence is variable in Anochetus , Bothroponera (present in members of the Bothroponera sulcata group), Brachyponera , Hypoponera , Mayaponera (only present in M. constricta ), Mesoponera , Myopias , Odontomachus , Phrynoponera (only present in P. pulchella ), and Ponera (Suppl. material 3: Table S3; see also Markl 1973; Schmidt and Shattuck 2014; Fisher and Bolton 2016). Schmidt and Shattuck (2014) state that it is universally present in Leptogenys and Platythyrea . However, Markl (1973), after a comprehensive taxa examination, affirmed that the occurrence of the trait is variable among species in both genera.

Contrary to Fisher and Bolton (2016), the stridulitrum is present in Brachyponera obscurans (Fig. 50A View Figure 50 ) and at least two Afrotropical and Malagasy species of Mesoponera (Fig. 50B, C View Figure 50 ). For the record, it is absent in Boloponera ikemkha . In addition, the midline of the fourth abdominal pretergite is strigulate and dissimilar to the surrounding sculpture in Euponera sikorae (Fig. 50D, E View Figure 50 ) and Mesoponera caffraria (Fig. 50F View Figure 50 ); the ridges are set farther apart and form a narrower area in the latter species. However, whether they have a stridulatory function in those species is unclear. Finally, the stridulitrum is present in males of Mayaponera pergandei (Fig. 50G View Figure 50 ; contrary to Mackay and Mackay 2010) but absent in the worker caste (Fig. 50H, I View Figure 50 ). Assuming that W. P. Mackay correctly determined the two males we examined, this intercaste variation is interesting because (1) it has not been observed in ants (see Markl 1973), and (2) it suggests that this trait has a reproductive function in M. pergandei .

47. The hypopygium (abdominal sternite VII) is armed with spine-like setae that flank the sting in some Ponerinae . According to previous studies, the setae are present in Dinoponera , Ophthalmopone , Pachycondyla , Paltothyreus , some Leptogenys , and few Ponera ( Schmidt and Shattuck 2014; Fisher and Bolton 2016). However, in Pachycondyla , we found that the hypopygial setae may be spine-like or aristate (Figs 10C-F View Figure 10 , 12C, D View Figure 12 ; see details in the preceding subsection "Transfers between Neoponera to Pachycondyla ").