Research Article |
Corresponding author: Denis J. Brothers ( brothers@ukzn.ac.za ) Academic editor: Michael Ohl
© 2017 Denis J. Brothers, Arkady S. Lelej.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Brothers DJ, Lelej AS (2017) Phylogeny and higher classification of Mutillidae (Hymenoptera) based on morphological reanalyses. Journal of Hymenoptera Research 60: 1-97. https://doi.org/10.3897/jhr.60.20091
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This study aimed to resolve the differences in the two currently used classifications of Mutillidae, which differ in many respects. Cladistic analyses of 101 genera and subgenera of Mutillidae (represented by females of 253 species and males of 260 species) and four outgroups (pepsine Pompilidae, anthoboscine Tiphiidae and both fedtschenkiine and sapygine Sapygidae) based on 230 morphological characters treated in various ways, produced most-parsimonious trees which were in broad agreement but differed in many details. Evaluation of these results led to the proposal of a compromise tree which reflected each proposed taxon as monophyletic, while trying to keep disruptions to the current classifications to a minimum. The result differs from both previous classifications, and proposes the recognition of eight subfamilies: Myrmosinae (with the tribes Kudakrumiini and Myrmosini), Pseudophotopsidinae, Rhopalomutillinae, Ticoplinae (with the tribes Smicromyrmillini and Ticoplini), Sphaeropthalminae (with the tribes Sphaeropthalmini, Dasymutillinitrib. n., and Pseudomethocini with the subtribes Euspinoliinasubtrib. n. and Pseudomethocina), Myrmillinae, Dasylabrinae (with the tribes Apteromutillinitrib. n. and Dasylabrini) and Mutillinae (with the tribes Ctenotillinitrib. n., Smicromyrmini, Mutillini with the subtribes Ephutina and Mutillina, and Trogaspidiini). Notably, Myrmosinae were consistently strongly supported as monophyletic with the remaining Mutillidae (disagreeing with a recent molecular analysis), and thus retained as a mutillid subfamily. The placements of all currently valid genera and subgenera in the proposed classification are provided.
Biogeography, cladistics, new tribe, new subtribe, parsimony, polymorphism, Sapygidae
The family Mutillidae (velvet-ants) includes approximately 4300 described species in 216 valid genera and 30 valid subgenera (
The higher classification of Mutillidae has changed considerably over time, but the first cladistic analysis of the aculeate Hymenoptera as a whole, by
Currently, there are thus two somewhat different classifications of Mutillidae being used (Fig.
Because of their extreme sexual dimorphism, we considered it essential to ensure that all terminals included in the study were known from both sexes, and preferably with at least one species represented by both sexes. We thus accumulated specimens of 101 sub/genera, including females of 253 species and males of 260 species of Mutillidae, and, as outgroups, we also included specimens of the three families which had previously been found to be those most closely related to Mutillidae in morphological analyses (
We scored all specimens individually for 230 characters derived from those previously considered by
Estimated phylogenies were derived under maximum parsimony using the Willi Hennig Society edition of TNT version 1.5 (
Several versions of the data were analysed, investigating the effects of additivity of character states, the influence of polymorphisms, and sexual differences: a) all terminals, considering all characters as non-additive/unordered; b) all terminals, considering many characters (those for which reasonable evolutionary sequences could be specified) as additive/ordered (as in Appendix
Since the results obtained for the various analyses differed in several respects, although generally reflecting a similar basic pattern, and it was not possible to determine which method was most likely to produce the “best” result, it was necessary to develop a compromise tree upon which the proposed classification could be based. Two basic principles were used in its construction. First, arrangements which would result in major disruptions to the currently used classifications were minimised, so as to promote nomenclatural stability as far as possible; this required a marked change in topology in only one instance. Second, paraphyletic groups for which the component terminals were separated by branches with only few and/or weak (homoplasious) apomorphies were rearranged so as to be reflected as monophyletic, also taking into account whether such groupings had been found to be supported by resampling in any of the analyses. Given the extent of homoplasy and polymorphism found for many of the characters, it was considered reasonable for the final compromise tree to be less than 1% longer than the comparable most parsimonious trees. Further details about the actual rearrangements proposed, and justifications for them, are provided below.
The initial analysis of all terminals based on both sexes was done employing minimal assumptions (all characters non-additive and equally weighted). The number of most-parsimonious trees (MPTs) found was 618 (length = 2633, ci = 0.20, ri = 0.59), and the strict consensus of these trees is shown in Fig.
Strict consensus of 618 most-parsimonious trees (length = 2633, ci = 0.20, ri = 0.59), of 101 sub/genera of Mutillidae and 4 outgroups, both sexes, 230 characters all non-additive and equally weighted. Group support (GC) values shown for all groups supported by resampling. Terminals in bold are those whose placements differ by more than mere taxonomic level in the classifications of DB and LN (see Appendix
Still considering all characters non-additive, the effect of implied weighting was then tested, and using N = 5 (k = 60), a single fully resolved tree was found which was only one step longer than the MPTs produced by the equal-weights analysis (raw length = 2634, ci = 0.20, ri = 0.59) (Fig.
Single most-parsimonious tree (raw length = 2634, ci = 0.20, ri = 0.59), of 101 sub/genera of Mutillidae and 4 outgroups, both sexes, 230 characters all non-additive and with implied weighting (N = 5, k = 60). Group support (GC) values shown for all groups supported by resampling. Terminals in bold are those whose placements differ by more than mere taxonomic level in the classifications of DB and LN (see Appendix
Analysis of all terminals for both sexes but considering all characters additive (except for those where a logical evolutionary sequence could not be postulated, see Appendix
Strict consensus of 38 most-parsimonious trees (length = 2828, ci = 0.19, ri = 0.61), of 101 sub/genera of Mutillidae and 4 outgroups, both sexes, 230 characters many additive and all equally weighted. Group support (GC) values shown for all groups supported by resampling. Terminals in bold are those whose placements differ by more than mere taxonomic level in the classifications of DB and LN (see Appendix
When implied weighting was applied (N = 5, k = 81), a single tree was found, one of the original MPTs (raw length = 2828, ci = 0.19, ri = 0.61) (Fig.
Single most-parsimonious tree (raw length = 2828, ci = 0.19, ri = 0.61), of 101 sub/genera of Mutillidae and 4 outgroups, both sexes, 230 characters many additive and all with implied weighting (N = 5, k = 81). Group support (GC) values shown for all groups supported by resampling. Terminals in bold are those whose placements differ by more than mere taxonomic level in the classifications of DB and LN (see Appendix
Analysis of the double-sized matrix (with duplicated and recoded terminals to explore polymorphisms, see above) under equal weights produced 98 MPTs (length = 3822, ci = 0.14, ri = 0.75), and implied weighting (N = 5, k = 169) found one of these trees. Fig.
Single most-parsimonious tree (raw length = 3822, ci = 0.14, ri = 0.75), of 101 sub/genera of Mutillidae and 4 outgroups (but each duplicated and recoded so as to reflect maximal character-state differences for polymorphisms, and taxa retained as monophyletic collapsed in the figure, see text), both sexes, 230 characters many additive and all with implied weighting (N = 5, k = 169). Group support (GC) values shown for all groups supported by resampling. Names in bold are of “terminals” shown not to be monophyletic.
When the reduced matrix (198 instead of 230 characters, those showing polymorphisms in at least 10% of the terminals having been deleted) was analysed under equal weights, 1330 MPTs (length = 2023, ci = 0.21, ri = 0.63) were found; using implied weighting (N = 5, k = 81), a single tree was found (length = 2024, ci = 0.21, ri = 0.63), only one step longer than the equal-weight MPTs (Fig.
Single most-parsimonious tree (raw length = 2024, ci = 0.21, ri = 0.63), of 101 sub/genera of Mutillidae and 4 outgroups both sexes, 198 characters (32 of the original 230 deleted, those found to be polymorphic in at least 10% of terminals) many additive and all with implied weighting (N = 5, k = 81). Group support (GC) values shown for all groups supported by resampling. Terminals in bold are those whose placements differ by more than mere taxonomic level in the classifications of DB and LN (see Appendix
In order to explore the degree to which the two sexes produced similar results (the tree/s found for each sex separately should at least be compatible and not contradictory if they actually are reflections of the evolutionary histories of the terminals) the characters of females and of males were analysed separately (seven characters applied to both sexes and so were included in both matrices). Analysis of the females (97 characters) considering all characters non-additive and of equal weight produced 358 MPTs (length = 1052, ci = 0.21, ri = 0.59), and under implied weighting (N = 5, k = 53) a single tree was found (raw length = 1057, ci = 0.20, ri = 0.59), five steps longer than the MPTs. When most characters were considered additive and all of equal weight 68 MPTs were found (length = 1131, ci = 0.19, ri = 0.61). Under implied weighting (N = 5, k = 81) a single tree was found (length = 1134, ci = 0.19, ri = 0.61), only three steps longer than the MPTs (Fig.
Single most-parsimonious tree (raw length = 1134, ci = 0.19, ri = 0.61), of 101 sub/genera of Mutillidae and 4 outgroups, females only, 97 characters many additive and all with implied weighting (N = 5, k = 81). Group support (GC) values shown for all groups supported by resampling. Terminals in bold are those whose placements differ by more than mere taxonomic level in the classifications of DB and LN (see Appendix
Single most-parsimonious tree (raw length = 1622, ci = 0.20, ri = 0.63), of 101 sub/genera of Mutillidae and 4 outgroups, males only, 140 characters many additive and all with implied weighting (N = 5, k = 71). Group support (GC) values shown for all groups supported by resampling. Terminals in bold are those whose placements differ by more than mere taxonomic level in the classifications of DB and LN (see Appendix
The results outlined above, as well as additional permutations which were tested, indicate that the structure near the base of the phylogeny is generally supported by a variety of analyses, and indicates a monophyletic Mutillidae, with the generally monophyletic subfamilies Myrmosinae, Pseudophotopsidinae, Ticoplinae and Rhopalomutillinae, but there is considerable variation in the groupings found above these taxa. Using the analysis of all terminals with additive characters of both sexes and implied weighting (identifying one of the MPTs as preferred) as the basis (see Figs
The arrangement of taxa in Fig.
The Myrmosinae has either included (DB) or excluded the Kudakrumiini/ae (LN). Either way, these two taxa have seemed clear-cut. The current analysis has shown, however, that the Kudakrumia–Myrmosula group is paraphyletic, with Myrmosula more closely related to Myrmosa–Paramyrmosa than to the other genera. Nevertheless, the arrangement shown necessitates that functional ocelli in the females were regained in Myrmosa–Paramyrmosa after having been lost in the ancestral mutillid (character 13, Fig.
The Pseudophotopsidinae includes only the genus Pseudophotopsis, but its species complicate the analysis because the females vary in having functional ocelli, reduced ocelli, or no ocelli whatsoever (character 13, Appendix
The Ticoplinae is clearly divided into the two accepted tribes, each comprising two terminals in this analysis and thus agrees with previous concepts.
The Sphaeropthalminae is clearly paraphyletic, with the Euspinolia–Hoplocrates group (considered as members of the pseudomethocine grouping by both DB and LN) arising as sister to the remainder of the Mutillidae. However, examination of Fig.
Liotilla, Brachymutilla and Apteromutilla appear as separate terminals sequentially diverging from the spine of the tree basal to the sphaeropthalmines; all three have completely apterous males with the mesosomal sutures entirely or substantially obliterated and both sexes very similar morphologically. They are restricted to southern Africa. Brachymutilla and Apteromutilla have previously been placed in the Dasylabrinae, but Liotilla (until now known only from the female holotype of its type species, L. laevis Bischoff) was placed in the Myrmillinae by Bischoff (1920). We have, however, recently been able to examine several species and both sexes of Liotilla, all collected in pitfall traps, which has enabled clarification of their relationships. When females only were analysed, these three genera appeared well separated on the tree (Fig.
The Myrmillinae (Viereckia–Platymyrmilla) formed a monophyletic group with low support in almost all of the analyses including both sexes (Figs
The Mutillinae (Pristomutilla–Trogaspidias.s.) formed a monophyletic group with low support in all of the analyses except for that of females only, which excluded Pristomutilla (Fig.
Although Fig.
The most contentious parts of the suggested rearrangements involve the Liotilla–Apteromutilla and Euspinolia–Hoplocrates groups, these together accounting for much of the increase in length of the tree. Their suggested placements, with Dasylabrinae and Pseudomethocini respectively, are not supported by resampling, however. Based on Fig.
Despite Fig.
Preferred most-parsimonious tree (see Fig.
Sapygidae + Mutillidae: As expected, the family Mutillidae is sister to Sapygidae, that association having good resampling support (here GC = 56), and supported by three unique and unambiguously placed synapomorphies for both additive and non-additive characters: 14.1 and 105.1, antennal “tubercle” in females and males (although further modified in male Sapyginae); 224.1, non-fusion of penis valves. An additional two unique and unambiguously placed synapomorphies were shown for the additive characters only: 118.2, short pleurostomal carina (although modified in Sapyginae); 209.1, posterior differentiation of sternum I in males (although further modified in Mutillidae).
Mutillidae Latreille, 1802: Monophyly of the Mutillidae (including Myrmosinae) has very high resampling support (GC = 99) and is supported by 10 unique and unambiguously placed synapomorphies for both additive and non-additive characters: 7.1, articulation of tergum II and sternum I in both sexes; 15.2, form of base of scape in females; 38.1, loss of wings in females; 65.2, closed metacoxal cavities in females; 90.1 and 208.1 stridulitrum on Tergum III in females and males (although apparently secondarily lost in some male myrmosines, and females of Rhopalomutillinae); 92.2 and 209.2, form of sternum I posteriorly in females and males; 200.1, reduction in jugal lobe of hind wing in males (although entirely lost subsequently); 225.2, form of penis valve (although subsequently modified in most terminals). There are also seven unambiguously placed but homoplasious states, the most significant being: 36.1, maxillary palp longer than fore tibia in females (but shorter in rhopalomutillines and Euspinolia, and even longer in scattered terminals); 61.3, metapleural-propodeal suture entirely obliterated on surface in females (but distinct in a few scattered terminals, and partially distinct in many; these apparently widespread reversals cast doubt on the accuracy of this placement); 127.1, maxillary palp longer than fore tibia in males (but shorter in Liotilla, and even longer in scattered terminals); 153.2, metapleural-propodeal suture obliterated dorsally and vague ventrally in males (but entirely distinct in a few scattered terminals, and partially distinct in many; these apparently widespread reversals cast doubt on the accuracy of this placement); 203.1, tergum I >0.5 <0.75 × width of tergum II in males (but broader in Hindustanilla and some Pseudophotopsis, even narrower in several scattered terminals). An additional unique and unambiguously placed state appears in the initially preferred tree (Fig.
Tree based on preferred tree (see Fig.
Myrmosinae Fox, 1894: This is a taxon whose estimated affinities have fluctuated in the past, and has recently been recognized again as a distinct family by
Kudakrumiini Krombein, 1979: In the originally preferred tree (Figs
Proposed higher classification of Mutillidae as reflected by the rearranged tree (Fig.
Myrmosini Fox, 1894: This group was found to be monophyletic with very high resampling support in all analyses (here GC = 97), and is supported by two unique and unambiguously placed synapomorphies for both additive and non-additive characters: 166.4, many meso- and metatibial articulated spines in males; 213.1, hypopygium concealed and modified in males. There are an additional nine unambiguously placed but homoplasious states supporting this, the most significant being: 79.1, tergum I with paired vertical ridges basally in females (found elsewhere only in some Ticoplinae); 228.0, basal lobe of volsella forming inner projection (found elsewhere only in some outgroups and a few scattered terminals). The tribe is Palaearctic, Nearctic and Oriental in distribution, with seven sub/genera; females are known for 71% and males for 100% of those taxa.
The remaining Mutillidae (apart from Myrmosinae) form a monophyletic group with very high resampling support in all analyses (here GC = 96), supported by four unique and unambiguously placed synapomorphies for both additive and non-additive characters: 4.2, metasternum with posterior median process(es) in both sexes; 79.2, tergum I with expanded “auricles” basally in females (although apparently reversed in a few Ticoplinae); 160.2, metacoxal cavities closed in males; 175.1, fore wing venation ending before distal margin of wing. There are an additional two unambiguously placed synapomorphies for additive characters only: 1.1, eye pubescence absent but pores present in both sexes (but subsequently modified in some groups); 42.1, pro-mesonotal suture distinct but fused in females (but subsequently modified in almost all). There are an additional 11 unambiguously placed but homoplasious states supporting the monophyly, the most significant being: 84.1, increased length of tergum II in females (subsequently modified in most terminals, reversed in Rimulotilla, and independently developed in Kudakrumia); 128.1, labial palp with mid segments expanded in males (although subsequently reversed in Liotilla, and independently developed in Myrmosa and one outgroup); 157.1, mid coxae slightly separated in males (but also in Kudakrumia and one outgroup); 194.1, hind wing crossvein r-m proximal (although subsequently modified in some); 205.1, felt line on tergum II present in males (although subsequently reversed in a few terminals; the female equivalent, 88.1, is not unambiguously placed here, but has a similar pattern).
Pseudophotopsidinae Bischoff, 1920: This group, comprising the single variable genus Pseudophotopsis, is confirmed as sister to the rest of the Mutillidae (except for Myrmosinae), in agreement with all previous analyses. Despite the small size of the group, it warrants recognition at the subfamily level, being morphologically very distinct, with a mixture of plesiomorphic (e.g., 13.0, presence of functional ocelli in females of some species; 42.1, distinct but fused pro-mesonotal suture in females; 189.0, 200.1, presence of a jugal lobe on both wings) and apomorphic states. It is supported by eight unique and unambiguously placed synapomorphies for both additive and non-additive characters: 2.1, pubescent pit on pronotum in both sexes; 6.1, pubescent depressions on sternum I in both sexes; 70.2, outer vertically elongate groove/pore on fore tibia in females; 136.4, interrupted faint parapsidal groove in winged males; 165.3, 167.3 and 173.2,3, pulvillus on 2nd–4th tarsomeres of all legs in males (absent on 2nd in some); 226.1, articulated spines on penis valve. It is also supported by several other unambiguously placed but homoplasious states (Fig.
The remaining Mutillidae (apart from Myrmosinae and Pseudophotopsidinae) also form a monophyletic group with very high resampling support in all analyses (here GC = 85), supported by seven unique and unambiguously placed synapomorphies for both additive and non-additive characters: 42.2, pro-mesonotal suture very indistinct or obliterated in females (although somewhat distinct in some Euspinolia species); 145.2, propodeal disc with three large fields in winged males (although apparently subsequently modified in most terminals since this state present only in Rhopalomutillinae and many Ticoplinae); 161.1, tarsal claws simple in males (subsequently modified in Rhopalomutillinae); 189.1 and 200.2, both wings without jugal lobe; 199.2, anal lobe not indicated on hind wing; 225.3, penis valve with simple apex and ventral tooth on apical half (but modified in Rhopalomutillinae). There are also several unambiguously placed but homoplasious states supporting this grouping (Fig.
Subtending states for tree reflecting proposed taxa as monophyletic. Blue indicates states found only when most characters were considered additive, red only when all states were considered non-additive, and black under both conditions. Solid hashmarks indicate unique state changes, and open hashmarks are homoplasies. Letters within boxes indicate breaks in branches to enable effective layout.
Ticoplinae Nagy, 1970: This group was found to be monophyletic with high resampling support (here GC = 73) in all analyses (except that for males only where it appeared as paraphyletic, Fig.
Smicromyrmillini Argaman, 1988: This group was found to be monophyletic with very high resampling support in all analyses (here GC = 99), and is supported by three unique but ambiguously placed synapomorphies for both additive and non-additive characters: 52.5, posterodorsal margin of propodeum with two median teeth and two lateral spines or teeth in females (but this present in only some species of Smicromyrmilla, so probably unreliable); 138.4, mesoscutellum posteriorly produced over metanotum in winged males; 215.4, hypopygium with complex narrow apical emargination (although plotted as ambiguous by Winclada, this state is unique to this group so is effectively unambiguously placed here). There are an additional eight unambiguously placed but homoplasious states supporting the group, the most significant being: 93.1, sternum II with felt line in females (although sporadically present in a few other terminals); 148.1, posterolateral margin of propodeum dentate or spinose in males (but also in Odontotillas.s. and some members of Ephutina); 217.5, cercus short, flattened basally and clavate apically in males (but also in most Rhopalomutillinae). In the analysis of duplicated terminals (Fig.
Ticoplini Nagy, 1970: This group was found to be monophyletic with moderate resampling support (here GC = 37) in all analyses except for that of males only (there paraphyletic, Fig.
The remaining Mutillidae (distal to Ticoplinae) also form a monophyletic group with good resampling support in all analyses (here GC = 62) , supported by five unique and unambiguously placed synapomorphies for both additive and non-additive characters: 60.1 and 152.1, meso-metapleural “bridge” present in females and males; 137.1, posterolateral margin of mesoscutum lobed in winged males (but sporadically subsequently modified or reversed in many terminals); 150.2, meso-metapleural suture fused in winged males; 190.1, basal hamuli on hind wing absent. The group is also supported by six unambiguously placed homoplasious synapomorphies, the most significant being: 28.2 and 121.2, oral and mandibular fossae separated by cuticular bridge in females and males (but reduced or elaborated in many subsequent terminals); 91.3, bounded pygidial plate present in females (but sporadically reduced or absent in many terminals); 134.1, mesoscutum extended far anterior to tegula in winged males (but sporadically shortened in several subsequent terminals).
Rhopalomutillinae Schuster, 1949: This group was found to be monophyletic with extraordinarily high resampling support in all analyses (here GC = 100), and is supported by four unique and unambiguously placed synapomorphies for both additive and non-additive characters: 35.2, maxillary palp unsegmented in females (although two-segmented in some species of Pherotilla and Rhopalomutilla): 75.1, metatibia broadened and smooth on inner surface in females; 161.2, tarsal claws lamellate basally and acute apically in males; 225.4, penis valve with rounded apex and ventral prominence at about half length. There are an additional 27 unambiguously placed but homoplasious states supporting the group, the most significant being: 20.0, flagellomere I wider than long in females (but also in Kudakrumia, Nanomutilla and Odontomyrme); 27.1 and 120.1, postmandibular carina present as blunt ridge in females and males (but also in Kudakrumia females, Lomachaeta and Liotilla); 36.0, maxillary palp shorter than protibia in females (but also in outgroups and Euspinolia); 40.2, mesosomal form in females (but similar in Protophotopsiss.s. and some Apteromutillina); 64.5, metasternal process long, unidentate and acute in females (but also in a few scattered Sphaeropthalminae); 80.1, tergum I posteriorly parallel-sided and discontinuous with tergum II in females (but similar in some Myrmosinae and Seyrigilla); 90.0, no stridulitrum on tergum III in females (but also in Paramyrmosa and Nanomutilla); 165.1, 167.1 and 173.1, pulvillus on 4th tarsomere of all legs in males (but also in various Mutillinae); 207.3, apical setae on tergum II strong and curved in males (but also in Protophotopsiss.s. and Darditilla, and straight with split apices in some Rimulotilla); 222.0, gonostylus without parapenial lobe (but also in some Dasylabrinae). The subfamily is Afrotropical and Oriental in distribution, with four genera; females and males are known for all genera. The genera were recently reviewed by
The remaining Mutillidae (distal to Rhopalomutillinae) form a poorly to moderately supported monophyletic group in all analyses (here GC = 44), but not supported by any unique and unambiguously placed synapomorphies for both additive and non-additive characters. There is a single unique but ambiguously placed synapomorphy: 202.2, tergum I gradually broadened, short and sessile posteriorly (but subsequently modified in various ways in many subsequent terminals). There are an additional 15 unambiguously placed but homoplasious states supporting the group, however, the most significant being: 5.0, metacoxa posterodorsally simple in both sexes (otherwise found only in some Sapyginae, so unique here in Mutillidae); 43.1, pronotum lateral length shorter than distance between prothoracic and propodeal spiracles in females (but also in Kudakrumia and Pherotilla, and reversed in a few Dasylabrinae and Mutillinae); 71.2 and 164.1, fore calcar blade expanded and longish in females and males (also only in Pseudophotopsidinae); 84.2, tergum II much longer than terga III–VI in females (but also in Nanomutilla, and about the same length in a few scattered subsequent terminals); 139.1, posterolateral surface of axilla concave in winged males (although subsequently modified in many terminals). The Liotilla–Apteromutilla and the Euspinolia–Hoplocrates groups appear in different positions in the proposed arrangement, as discussed above.
Sphaeropthalminae Schuster, 1949 (1903): This group was paraphyletic in most of the analyses, with the Euspinolia–Hoplocrates group appearing as sister to the group containing the rest of the Sphaeropthalminae and the remaining Mutillidae, but this with negligible or no resampling support (Figs
Sphaeropthalmini Schuster, 1949 (1903): This group comprises those sphaeropthalmines which are more basal than those more closely related to the pseudomethocines (see above). The Cystomutilla–Scaptodactyla group is moderately well supported (here GC = 39) in all of the analyses (except that of females only), but the positions of Tallium, Allotilla and Photomorphuss.s. vary somewhat. Using Fig.
The remaining Sphaeropthalminae (distal to Sphaeropthalmini, and disregarding the Euspinolia–Hoplocrates group) form a poorly supported monophyletic group in most analyses, but the proposed combined group is not supported by resampling or by any unique and unambiguously placed synapomorphies. There is, however, a single unique but ambiguously placed synapomorphy for both additive and non-additive characters: 102.6, eye subcircular with convex inner margin and long axis horizontal in males (but long axis vertical in the Euspinolia–Hoplocrates group and some Dimorphomutilla). It is also supported by three unambiguously placed homoplasious synapomorphies: 21.1, head with genal carina in females (but also in several other terminals, and absent in a few); 101.2, eye strongly convex in males (but also in some Sphaeropthalmini and Tricholabiodes, and only moderately convex in Euspinolia and Myrmilloides); 145.0, propodeal disc evenly sculptured in winged males (but also in a few Sphaeropthalmini, some Dasylabrinae, a few Mutillinae and many basal-most mutillids, and different in Bothriomutilla, Euspinolia and Vianatilla).
Dasymutilla Ashmead, 1899. This group is paraphyletic in most analyses, although, interestingly, monophyletic in the tree derived from males only (Fig.
Pseudomethocini Brothers, 1975: The two components of this grouping are not closely associated in any of the analyses, but they are placed together here on the basis of their consistent positions in the current classifications, and the fact that this arrangement adds only five steps when compared with that in the preferred tree (Fig.
Euspinolia Ashmead, 1903. This group was found to be monophyletic in all analyses with very high resampling support (here GC = 90), but somewhat inconsistent in its placement; our justification for including it in the Pseudomethocini appears above. The close association of Atillum and Hoplocrates has long been recognized, but the inclusion of Euspinolia with them and separate from the other pseudomethocines is unexpected. The group is not supported by any unique and unambiguously placed synapomorphies, but there is one unique but ambiguously placed synapomorphy for both additive and non-additive characters: 163.2, fore tibia with obliquely elongate outer secretory pore in males (but absent in some Euspinolia). There are 13 unambiguously placed homoplasious synapomorphies, the most significant being: 70.3, fore tibia with obliquely elongate outer secretory pore in females (also only in Ronisia); 82.0 and 201.0, tergum I and/or propodeum with simple pubescence in females and males (within Sphaeropthalminae also only in Cephalomutilla, Gogoltilla, some Dasymutilla, some Bothriomutilla females, and Lophomutilla males); 99.0, head with simple pubescence in males (within Sphaeropthalminae also only in Cephalomutilla); 219.1 and 220.1, gonostylus (paramere) short, tapered and apically straight (within Sphaeropthalminae also only in Myrmilloides). The subtribe is Neotropical, with three genera; females and males are known for all genera.
Pseudomethocina Brothers, 1975: This group was found to be monophyletic in all analyses (except for that of females only) with low to moderate resampling support (here GC = 39). The group is not supported by any unique synapomorphies, but there are three ambiguously placed homoplasious synapomorphies for both additive and non-additive characters: 12.1, ommatidia faintly distinguishable in females (but also in several terminals elsewhere and further modified in some here); 73.1, metacoxa carinate mesad in females (but also in several other groups); 140.2, axilla anterolaterally with broad vertical flange in winged males (but also in several Sphaeropthalminis.s.). Within the group, the position of Pseudomethoca differs from that in the preferred tree (Fig.
The remaining Mutillidae (distal to Sphaeropthalminae, and disregarding the proposed inclusion of the Liotilla–Apteromutilla group) form a monophyletic group in the preferred tree (Fig.
Dasylabrinae Invrea, 1964: This group (disregarding the Liotilla–Apteromutilla group which is now placed here as a distinct tribe) was found to be monophyletic in some of the analyses, including the preferred tree (Figs
Apteromutilla Ashmead, 1903. Although the terminals in this group were closely associated in most analyses (see above), and it has low resampling support here (GC = 19), it is not supported by any unique synapomorphies, but there are six unambiguously placed homoplasious synapomorphies for both additive and non-additive characters, the most significant being: 40.2, mesosomal form in females (also in rhopalomutillines and Protophotopsiss.s., and modified in Liotilla); 110.2, pedicel distinctly longer than wide in males (also in Hindustanilla only); 131.1, humeral angle blunt in males (also in some scattered terminals, and carinate in some Liotilla); 174.3, apterous without any trace of wings or tegula in males (also only in Hindustanilla and some Viereckia); 219.1, gonostylus (paramere) short and narrow (also in ticoplines, some myrmosines and sphaeropthalmines, Dasylabroides and Dasylabris, and lamellate in Brachymutilla). Of interest is that Brachymutilla and Liotilla are apparently the only Mutillidae to lack cerci in the males (Fig.
Dasylabrini Invrea, 1964: Although a Stenomutilla group was separated from a Dasylabris group in several analyses (see above), the position of Chrestomutilla varied, being associated with either group. There is thus no good reason to recognize these subgroups formally. The group is not supported by resampling nor by any unique synapomorphies, but there are six unambiguously placed homoplasious synapomorphies for both additive and non-additive characters, the most significant being: 49.1, mesosoma with scutellar scale in females (also in most smicromyrmines, many trogaspidiines and several other scattered terminals, and absent in some here); 202.1, tergum I >0.5 × length of tergum II and apically constricted in males (also in most rhopalomutillines and sphaeropthalmines s.s. and a few scattered terminals, tergum I shorter in Dasylabris and Chrestomutilla); 220.0, gonostylus (paramere) apically upcurved (also in most sphaeropthalmines and a few other scattered terminals). In the analysis of duplicated terminals (Fig.
The remaining Mutillidae (distal to Dasylabrinae) form a monophyletic group with low to moderate resampling support (here GC = 34) in all of the analyses (except for that of females only, which associated the Euspinolia–Hoplocrates group here, and in which most groupings had no resampling support). The group is not supported by any unique synapomorphies, but there are 10 ambiguously placed homoplasious synapomorphies for both additive and non-additive characters, the most significant being: 40.0, mesosoma parallel-sided in females (also in Areotilla only, but subsequently modified in a few scattered terminals); 58.1, meso-metapleural suture strongly angled in females (also only in Tallium, Darditilla and some Lophomutilla, and weakly curved in several myrmillines); 179.1, fore wing with constriction in Sc+R only at pterostigmal base (also in several scattered terminals and many dasymutillines, and subsequently modified in some terminals here); 188.1, fore wing crossvein 3r-m without bulla (also in rhopalomutillines, and with bulla in mutillines s.s. and Dolichomutilla). There are also two ambiguously placed homoplasious synapomorphies for the analysis using additive characters: 54.2, mesopleuron with dorsal region depressed in females (also in ticoplines, some pseudomethocines, Kudakrumia and Pseudophotopsis, subsequently modified in a few scattered terminals here); 153.1, metapleural-propodeal pleural suture obliterated dorsally but distinct ventrally in winged males (also in many dasymutillines and a few other terminals, and modified in a few terminals here).
Myrmillinae Bischoff, 1920: This group was found to be monophyletic with slight to low resampling support (here GC = 23) in most analyses (excepting only the unweighted analysis using additive characters, consensus tree in Fig.
Mutillinae Latreille, 1802: This group was found to be monophyletic with low resampling support (here GC = 20) in most analyses (excepting that of females only (Fig.
Ctenotilla Bischoff, 1920. A group including four terminals (Mimecomutillas.s.–Ctenotilla) was found to be monophyletic in all analyses with high resampling support and almost always with Pristomutilla just basal to it, although generally without support; Pristomutilla was more distant in the analysis in which the most-polymorphic characters had been deleted (Fig.
The remaining Mutillinae (distal to Ctenotillini) formed a monophyletic group with low resampling support in all of the analyses (here GC = 15), except for that where the most-polymorphic characters were deleted (Fig.
Smicromyrmini Bischoff, 1920: This group was found to be monophyletic with some resampling support only in the weighted analysis of non-additive characters (Fig.
The remaining Mutillinae (distal to Smicromyrmini) formed a monophyletic group, although with no or negligible resampling support, in most of the analyses, except for the equal-weights analysis of non-additive characters (Fig.
Mutillini Latreille, 1802: This group was found to be monophyletic, although with weak to no resampling support, in both analyses of additive characters (Figs
Ephutina Ashmead, 1903: This group corresponds to the Ephutinae of the LN classification. The association of Ephuta and Odontomutilla/Yamanetilla is intuitively surprising, since they appear very different morphologically, but it is strongly supported in all analyses (here resampling support is very high, GC = 91). The group is supported by one unique and unambiguously placed synapomorphy for both additive and non-additive characters: 117.3, hypostomal carina strong anterolaterally but obsolete posteriorly in males (not found elsewhere); there is also one unique but ambiguously placed synapomorphy: 89.1, tergum II with felt line a broad patch in females (not found elsewhere, but subsequently uniquely modified in Odontomutilla and Yamanetilla, so questionably a synapomorphy here). In addition there are 15 unambiguously placed homoplasious synapomorphies, the most significant being: 23.2, gena with strong tooth anteroventrally in females (also in Atillum and Pertyella, and absent in some Odontomutilla); 24.4, hypostomal carina strong anterolaterally but obsolete posteriorly in females (also in Scaptodactyla and Radoszkowskitilla); 26.2 and 119.2, postgenal ridge distinct and merging with hypostomal carina in females and males (also in females of Bothriomutilla, Odontomyrme and some Mimecotilla, and in males of Rhopalomutilla); 73.0, metacoxa smoothly rounded mesad (also in many “more-basal” terminals but in no other mutillines); 118.3, pleurostomal carina long and straight with hypostomal carina to outer mandibular articulation (also in some scattered terminals elsewhere but in no other mutillines); 178.1, fore wing with vein SC lost or much reduced and pterostigma not delimited basally (also in most myrmillines and a few scattered terminals elsewhere but in no other mutillines). In the analysis of duplicated terminals (Fig.
Mutillina Latreille, 1802: This group was monophyletic (with low resampling support) only in the analyses using non-additive characters (Figs
Trogaspidiini Bischoff, 1920: Disregarding Dolichomutilla (sometimes associated with Mutillini), this group was found to be monophyletic in most analyses: those of non-additive characters (Figs
The variations seen in the results of the different analyses of this most-representative sample of Mutillidae examined to date, including both sexes, many more characters than previous efforts, and aspects of polymorphism, cast doubt on the accuracy of any one of the approaches to be a best estimate of the actual phylogeny/evolutionary history of the components of the family. It is also evident that including many more exemplars and characters, and not using groundplans, has greatly complicated the results, but probably made them more realistic. Consequently, we have proposed a compromise higher classification which takes the results of our various analyses into account and amalgamates them, and also deviates from the current classifications as little as possible, but thereby provides an informed framework for future studies (Fig.
Over many years, numerous curators and colleagues have donated or lent specimens to both authors from their own or institutional collections, without which this investigation would not have been possible; we express our extreme gratitude to all of them for their interest, generosity and patience. Kevin Williams kindly supplied information associating the sexes of Cephalomutilla through molecular-genetic analyses, thus enabling us to include the genus in our study. We are also grateful to the reviewers, Wojciech Pulawski and Gavin Broad, for their comments which improved the manuscript, and to Michael Ohl for his editorial assistance. This work was partially supported by funding from the Russian Foundation of Basic Research (# 15–29–02466, # 17–04–00259) for ASL, and the Research Office of the University of KwaZulu-Natal for DJB.
Specimens examined for phylogenetic analysis of sub/genera of Mutillidae and four outgroup taxa.* = type species of relevant genus/subgenus; “Spp. ♀♀” and “Spp. ♂♂” = number of species represented by female and male specimens respectively; “% poly.” = proportion of characters showing polymorphisms in taxon, values above 9% in bold; “Current classification” = placement as in specified papers, or as appropriate for taxa described later (see Fig.
Taxon | Spp. ♀♀ | Spp. ♂♂ | % poly. | Current classification | |
---|---|---|---|---|---|
DB | LN | ||||
Pompilidae, Pepsinae: Hemipepsis Dahlbom, 1843: H. capensis (Linnaeus, 1764) ♀, ♂, South Africa; *H. errabunda (de Dalla Torre, 1897) ♀, ♂, South Africa; H. ? hilaris (Smith, 1879) ♀, ♂, South Africa | 3 | 3 | 1% | - | - |
Tiphiidae, Anthoboscinae: Anthobosca Guérin de Méneville, 1838: A. spp. ♀♀, ♂♂, South Africa | 2 | 2 | 1% | - | - |
Sapygidae, Fedtschenkiinae: Fedtschenkia de Saussure, 1880: F. grossa de Saussure, 1880 ♀, ♂, Turkmenistan; F. anthracina (Ashmead, 1898) ♀, ♂, USA | 2 | 2 | 1% | - | - |
Sapygidae, Sapyginae: Krombeinopyga pumila (Cresson, 1880) ♀, ♂, USA; Polochrum sp. ♀, USA; Sapygina sp. ♂, South Africa | 2 | 2 | 7% | - | - |
Acanthomutilla Nonveiller, 1995: *A. comparanda (Bischoff, 1920) ♀, ♂, Zambia, Zimbabwe | 1 | 1 | 0% | Smicromyrmina | Myrmillinae |
Allotilla Schuster, 1949: *A. gibbosa Schuster, 1949 ♀, ♂, Paraguay | 1 | 1 | 0% | Pseudomethocina | Pseudomethocini |
Amblotropidia Nonveiller, 1995: *A. aurea (Bischoff, 1920) ♂, Cameroon; A. niveomaculata (André, 1898) ♀, ♂, Eritrea | 1 | 2 | 5% | Smicromyrmina 1 | Trogaspidiini 1 |
Ancistrotilla Brothers, 2012: A. aenigmatica Brothers, 2012 ♂, New Caledonia; *A. azurea Brothers, 2012 ♂, Vanuatu; A. caledonica (André, 1896) ♀, New Caledonia; A. ? depressa (Smith, 1879) ♀, ♂, Australia; A. fabricii (André, 1898) ♀, ♂, Australia | 3 | 4 | 2% | Sphaeropthalminae | Sphaeropthalmini |
Apteromutilla Ashmead, 1903: *A. aede (Péringuey, 1899) ♂, South Africa; A. aethra (Péringuey, 1899) ♀, South Africa; A. spp. ♀♀, ♂♂, South Africa | 3 | 3 | 1% | Dasylabrini | Dasylabrinae |
Areotilla Bischoff, 1920: *A. areolata Bischoff, 1920 ♂, Lesotho; A. marshalli (André, 1903) ♂, Malawi; A. perplexa Mitchell & Brothers, 1998 ♀, South Africa; A. vulgaris Mitchell & Brothers, 1998 ♂, South Africa | 1 | 3 | 1% | Ticoplini | Ticoplini |
Artiotilla Invrea, 1950: *A. biguttata (Costa, 1858) ♀, ♂, Cyprus, Montenegro | 1 | 1 | 0% | Smicromyrmina 1 | Petersenidiini 1 |
Atillum André, 1902: A. albicomum Mickel, 1943 ♀, Argentina; A. allophylum Mickel, 1943 ♀, Argentina; A. jucundum Mickel, 1943 ♀, Argentina; A. picturatum Mickel, 1943 ♂, Argentina; A. nr. optabile Mickel, 1943 ♂♂, Argentina; A. nr. picturatum Mickel, 1943 ♂, Argentina | 3 | 4 | 4% | Pseudomethocina | Pseudomethocini |
Bischoffiella Brothers, 2015: *B. cristata (Bingham, 1912) ♀, ♂, Zimbabwe; B. sp. ♀, ♂, South Africa | 2 | 2 | 1% | Rhopalomutillinae | |
Bischoffitilla Lelej, 2002: B. byblis (Mickel, 1934) ♀, Philippines; B. clypealis (Mickel, 1935) ♂, Malaysia; B. spp. ♀, ♂♂, India, Malaysia, Vietnam | 2 | 3 | 5% | Myrmillinae | Myrmillinae |
Bothriomutilla Ashmead, 1899: *B. rugicollis (Westwood, 1843) ♀, ♂, Australia | 1 | 1 | 0% | Sphaeropthalminae | Sphaeropthalmini |
Brachymutilla André, 1901: *B. androgyna (André, 1901) ♂, South Africa; B. gynandromorpha (André, 1901) ♂, South Africa; B. namana Bischoff, 1920 ♀, Namibia; B. peringueyi Bischoff, 1920 ♀, South Africa; B. scabrosa Bischoff, 1920 ♀, ♂, South Africa; B. spp. ♀, ♂, Namibia | 4 | 4 | 6% | Dasylabrini | Dasylabrinae |
Cephalomutilla André, 1908: C. ? confluenta Mickel, 1960 ♀, Argentina; *C. graviceps (André, 1903) ♀, Argentina; C. ? vulnerifera (André, 1908) comb. n. ♂, Argentina; C. nr. vulnerifera (André, 1908) ♂, Argentina | 2 | 2 | 1% | Sphaeropthalminae | Sphaeropthalmini |
Ceratotilla Bischoff, 1920: *C. dolosa (Smith, 1879) ♀, ♂, South Africa; C. spp. ♀♀, South Africa | 3 | 1 | 4% | Myrmillinae | Myrmillinae |
Chaetomutilla Nonveiller, 1979: *C. fornasinii (Gribodo, 1894) ♀, ♂, South Africa; C. lobognatha (André, 1902) ♂, South Africa | 1 | 2 | 2% | Smicromyrmina | Mutillini |
Chrestomutilla Brothers, 1971: C. ? maja (Péringuey, 1898) ♀, ♂, South Africa | 1 | 1 | 0% | Dasylabrini | Dasylabrinae |
Ctenotilla Bischoff, 1920: *C. caeca (Radoszkowski, 1879) ♀, ♂, Armenia, Crimea; C. guangdongensis Lelej, 1992 ♀, ♂, China, Laos | 2 | 2 | 1% | Smicromyrmina | Mutillini |
Cystomutilla André, 1896: *C. ruficeps (Smith, 1855) ♀, ♂, Croatia, France; C. teranishii Mickel, 1935 ♀, ♂, Japan | 2 | 2 | 0% | Sphaeropthalminae | Sphaeropthalmini |
Darditilla Casal, 1965: D. araxa (Cresson, 1902) ♀, Paraguay; D. garciai Casal, 1968 ♀, Argentina; D. spp. ♂♂, Brazil, Costa Rica | 2 | 2 | 3% | Pseudomethocina | Pseudomethocini |
Dasylabris Radoszkowski, 1885: D. m. maura (Linnaeus, 1758) ♀, ♂, France; D. maura sungora (Pallas, 1773) ♀, Kazakhstan; D. mephitis (Smith, 1855) ♀, ♂, South Africa; D. siberica (Christ, 1791) ♂, Russia; D. stimulatrix (Smith, 1879) ♀, ♂, South Africa | 3 | 5 | 5% | Dasylabrini | Dasylabrinae |
Dasylabroides André, 1901: D. bechuana Péringuey, 1914 ♂, Namibia; D. caffra (Kohl, 1882) ♀, South Africa; D. canace (Péringuey, 1899) ♀, South Africa; *D. capensis (Saussure, 1867) ♀, South Africa; D. ? neavei André, 1909 ♀, Zambia; D. phylira (Péringuey, 1898) ♂, South Africa; D. nr. idia (Péringuey, 1899) ♀, ♂, South Africa | 4 | 4 | 11% | Dasylabrini | Dasylabrinae |
Dasymutilla Ashmead, 1899: D. dilucida Mickel, 1928 ♀, USA; *D. gorgon (Blake, 1871) ♀, USA; D. melancholica (Smith, 1879) ♀, ♂, Dominican Republic; D. occidentalis (Linnaeus, 1758) ♀, ♂, USA; D. quadriguttata (Say, 1823) ♀, ♂, USA; D. vestita (Lepeletier, 1845) ♂, USA | 5 | 4 | 18% | Sphaeropthalminae | Sphaeropthalmini |
Dentilla Lelej, 1980: D. dichroa (Sichel & Radoszkowski, 1869) ♂, Afghanistan; *D. curtiventris (André, 1901) ♀, ♂, Armenia; D. persica (Sichel & Radoszkowski, 1869) ♀, ♂, Armenia, Greece; D. saharica (Giner Mari, 1945) ♀, ♂ Algeria, Morocco, Tunisia | 3 | 4 | 4% | Smicromyrmina | Smicromyrmini |
Dilophotopsis Schuster, 1958: *D. concolor (Cresson, 1865) ♂, Mexico, USA; D. stenognatha Schuster, 1958 ♀, ♂, USA | 1 | 2 | 3% | Sphaeropthalminae | Sphaeropthalmini |
Dimorphomutilla Ashmead, 1903: D. formosa Mickel, 1938 ♀, Chile; D. herbsti (André, 1904) ♂, Chile; D. ? punctifera Mickel, 1938 ♂, Chile; D. reedi Mickel, 1938 ♀, Chile; D. suavissima (Gerstaecker, 1874) ♀, ♂, Chile | 3 | 3 | 5% | Pseudomethocina | Pseudomethocini |
Dolichomutilla Ashmead, 1899: D. conigera (André, 1896) ♂, Cameroon; D. livingstonis (Kohl, 1882) ♀, South Africa; D. minor minor Bischoff, 1920 ♀, ♂, South Africa; D. scutellifera (André, 1894) ♀, Cameroon; D. sycorax (Smith, 1855) ♀, ♂, South Africa | 4 | 3 | 4% | Smicromyrmina 1 | Trogaspidiini 1 |
Ephuta Say, 1836: E. ? arpala Casal, 1968 ♂, Brazil; E. ? huavunca Casal, 1968 ♀, Argentina; E. s. sabaliana Schuster, 1951 ♂, USA; E. nr. aillanca Casal, 1968 ♀, Argentina; E. nr. melina Casal, 1968 or sauca Casal, 1968 ♀, Argentina; E. ? spinifera Schuster, 1951 ♀, USA; E. spp. ♀♀, ♂, Mexico, Panama; E. ? tapiola Casal, 1968 ♂, Argentina | 6 | 4 | 12% | Ephutini 2 | Ephutini 2 |
Ephutomma Ashmead, 1899: E. angustata (Skorikov, 1935) ♀, ♂, Kazakhstan, Turkmenistan; *E. turcestanica (de Dalla Torre, 1897) ♀, ♂, Kazakhstan, Turkmenistan | 2 | 2 | 0% | Smicromyrmina | Smicromyrmini |
Eurygnathilla Skorikov, 1927: *Myrmilla (E.) ephutommatina Skorikov, 1927 ♀, ♂, Turkmenistan, Uzbekistan | 1 | 1 | 0% | Myrmillinae | Myrmillinae |
Eurymutilla Ashmead, 1899 (near): nr. E. spp. ♀♀♀, ♂♂, Australia | 3 | 2 | 1% | Sphaeropthalminae | Sphaeropthalmini |
Euspinolia Ashmead, 1903: E. ? albicoma Mickel, 1938 ♂, Chile; E. canescens Mickel, 1938 ♂, Chile; E. clypeata Mickel, 1938 ♀, Chile; E. insignita Mickel, 1938 ♀, Chile; E. irregularis (Smith, 1879) ♂, Chile; E. militaris Mickel, 1938 ♀, Chile | 3 | 3 | 6% | Pseudomethocina | Pseudomethocini |
Glossotilla Bischoff, 1920: G. adelpha fuelleborni Bischoff, 1920 ♀, ♂, South Africa; G. suavis speculatrix (Smith, 1879) ♀, ♂, South Africa | 2 | 2 | 1% | Smicromyrmina 1 | Trogaspidiini 1 |
Gogoltilla Williams, Brothers & Pitts, 2011: *G. chichikovi Williams, Brothers & Pitts, 2011 ♀, ♂ | 1 | 1 | 0% | Pseudomethocina | Pseudomethocini |
Hemutilla Lelej, Tu & Chen in Tu et al., 2014: H. bifurcata (Chen, 1957) ♀, China; H. cheni Tu & Lelej in Tu, Lelej & Chen, 2014 ♀, China; H. ferrugineipes Tu, Lelej & Chen, 2014 ♂, China; *H. granulata Tu, Lelej & Chen, 2014 ♂, China; H. hoozana (Zavattari, 1913) ♂, China; H. tuberculata Tu, Lelej & Chen, 2014 ♂, China | 2 | 4 | 6% | Sphaeropthalminae | Sphaeropthalmini |
Hindustanilla Lelej in Lelej & Krombein, 2001: *H. indica Lelej in Lelej & Krombein, 2001 ♂, India; H. nathani Lelej in Lelej & Krombein, 2001 ♀, India; H. sp. ♂, India | 1 | 2 | 1% | Smicromyrmillini | Smicromyrmillini |
Hoplocrates Mickel, 1937: *H. cephalotes (Swederus, 1787) ♀, Brazil; H. ? mystica (Gerstaecker, 1874) ♂, Brazil; H. pompalis Mickel, 1941 ♀, Trinidad; H. speculatrix (Gerstaecker, 1874) ♀, ♂, Brazil; H. tartarina Mickel, 1941 ♀, Ecuador | 4 | 2 | 7% | Pseudomethocina | Pseudomethocini |
Hoplognathoca Suárez, 1962: H. costarricensis Suárez, 1962 ♀, ♂, Costa Rica | 1 | 1 | 0% | Pseudomethocina | Pseudomethocini |
Hoplomutilla Ashmead, 1899: H. acutangula (Gerstaecker, 1847) ♂, Venezuela; H. caerulea Mickel, 1939 ♂, Venezuela; H. gigantea (Perty, 1833) ♀, Brazil; H. opima Mickel, 1939 ♀, ♂, Trinidad; H. panamensis Mickel, 1939 ♀, Panama; H. rapax Mickel, 1939 ♀, Ecuador | 4 | 3 | 5% | Pseudomethocina | Pseudomethocini |
Karlissaidia Lelej, 2005: *K. medvedevi Lelej, 2005 ♀, ♂, Sri Lanka; K. turneri Lelej, 2005 ♀, Sri Lanka; K. sexmaculata (Swederus, 1787) comb. n. ♀, ♂, India | 3 | 2 | 12% | Smicromyrmina 1 | Trogaspidiini 1 |
Krombeinidia Lelej, 1996: K. lilliputiana (André, 1894) ♂, India; *K. peterseni Lelej, 1996 ♀, ♂, Sri Lanka; K. sp. ♀, Sri Lanka | 2 | 2 | 2% | Smicromyrmina 1 | Petersenidiini 1 |
Kudakrumia Krombein, 1979: *K. mirabilis Krombein, 1979 ♀, ♂, Sri Lanka | 1 | 1 | 0% | Kudakrumiini | Kudakrumiinae |
Labidomilla André, 1902: L. subinermis André, 1903 ♀, South Africa; *L. tauriceps (Kohl, 1882) ♀, ♂, South Africa; L. spp. ♀♀, ♂♂♂, Malawi, South Africa | 4 | 4 | 10% | Myrmillinae | Myrmillinae |
Liotilla Bischoff, 1920: L. spp. ♀♀♀♀, ♂♂♂♂♂, Botswana, Namibia, South Africa | 4 | 5 | 3% | Myrmillinae | Myrmillinae |
Lobotilla Bischoff, 1920: *L. leucopyga (Klug, 1829) ♀, ♂, Cameroon; L. leucospila (Cameron, 1910) ♀, ♂, South Africa | 2 | 2 | 1% | Smicromyrmina 1 | Trogaspidiini 1 |
Lomachaeta Mickel, 1936: *L. hicksi Mickel, 1936 ♀, ♂, USA | 1 | 1 | 0% | Sphaeropthalminae | Sphaeropthalmini |
Lophomutilla Mickel, 1952: L. prionophora (Burmeister, 1866) ♀, Brazil; L. seabrai Casal, 1968 ♀, Brazil; L. spp. ♂♂, Brazil, Costa Rica | 2 | 2 | 4% | Sphaeropthalminae | Sphaeropthalmini |
Lynchiatilla Casal, 1963: L. hoplites (Gerstaecker, 1874) ♀, Argentina; L. leguera Casal, 1963 ♀, ♂, Argentina; L. ? chayera Casal, 1963 ♂, Argentina; L. tacana Casal, 1963 ♀, Argentina | 3 | 2 | 2% | Pseudomethocina | Pseudomethocini |
Mickelomyrme Lelej, 1995: M. ? exacta (Smith, 1879) ♂, Laos; *M. hageni (Zavattari, 1913) ♀, ♂, Japan; M. ? kuznetsovi Lelej, 1996 ♀, Laos; M. yunnanensis Lelej, 1996 ♂, Laos | 2 | 3 | 6% | Smicromyrmina | Smicromyrmini |
Mimecomutilla Ashmead, 1903: M. (M.) renominanda Bischoff, 1920 ♀, ♂, South Africa; M. (M.) umtata (Péringuey, 1909) ♀, ♂, South Africa | 2 | 2 | 2% | Smicromyrmina | Mutillini |
Mimecotilla Nonveiller, 1998: Mimecomutilla (M.) bitaeniata Bischoff, 1920 ♀, ♂, South Africa; *Mimecomutilla (M.) nyassicola Bischoff, 1920 ♀, ♂, Cameroon | 2 | 2 | 4% | Smicromyrmina | Mutillini |
Mutilla Linnaeus, 1758: M. coerulea Bischoff, 1920 ♂, Cameroon; *M. europaea Linnaeus, 1758 ♀, ♂, Austria, Bosnia, Switzerland; M. quinquemaculata Cyrillo, 1797 ♀, ♂, Cyprus, Malta; M. scabrofoveolata penicillata André, 1895 ♀, South Africa | 3 | 3 | 7% | Mutillina | Mutillini |
Myrmilla Wesmael, 1851: *M. calva (Villers, 1789) ♀, ♂, Greece, Serbia, Spain; M. erythrocephala (Latreille, 1792) ♀, ♂, Cyprus, Greece | 2 | 2 | 3% | Myrmillinae | Myrmillinae |
Myrmilloides André, 1902: *M. grandiceps (Blake, 1872) ♀, ♂, USA | 1 | 1 | 0% | Pseudomethocina | Pseudomethocini |
Myrmosa Latreille, 1797: *M. atra Panzer, 1801 ♀, ♂, Denmark, Italy; M. unicolor Say, 1824 ♀, ♂, USA | 2 | 2 | 2% | Myrmosini | Myrmosinae |
Myrmosula Bradley, 1917: *M. parvula (Fox, 1893) ♀, ♂, USA; M. rutilans (Blake, 1879) ♀, USA; M. nr. rufiventris (Blake, 1879) ♂, USA | 2 | 2 | 0% | Kudakrumiini | Kudakrumiinae |
Nanomutilla André, 1900: *N. vaucheri (Tournier, 1895) ♀, Gibraltar; N. spp. ♀♀, ♂♂♂, South Africa, Zimbabwe | 3 | 3 | 3% | Ticoplini | Ticoplini |
Nemka Lelej, 1985: N. viduata bartholomaei (Radoszkowski, 1865) ♀, ♂, Kazakhstan; N. viduata insulae (Invrea, 1940) ♀, ♂, Cyprus; *N. v. viduata (Pallas, 1773) ♀, ♂, Czech Republic, Greece, Italy, Slovakia | 3 | 3 | 1% | Smicromyrmina | Smicromyrmini |
Odontomutilla Ashmead, 1899: O. ? aegrota (Cameron, 1898) ♀, Zimbabwe; O. ? chione (Péringuey, 1898) ♀, Lesotho, South Africa; O. ? chionella Bischoff, 1920 ♂, Lesotho; O. ? cleopatra (Péringuey, 1899) ♂, South Africa; O. ? fracta (Saussure, 1891) ♀, Kenya; O. ? inanis Mickel, 1935 ♀, Papua New Guinea; O. pulchrina (Smith, 1855) ♀, ♂, India; O. nr. calida André, 1908 ♀, Zambia; O. nr. tamensis (Cameron, 1906) ♀, Australia; O. tisiphonella Bischoff, 1920 ♂, South Africa; O. ? tomyris (Péringuey, 1899) ♀, South Africa | 8 | 4 | 11% | Mutillina | Odontomutillini |
Odontomyrme Lelej, 1983: O. spp. ♀♀♀♀♀, ♂♂♂, Australia, Papua New Guinea | 5 | 3 | 2% | Sphaeropthalminae | Odontomutillini |
Odontophotopsis Viereck, 1903: O. inconspicua (Blake, 1886) ♀, ♂, USA; O. villosa Mickel in Mickel & Clausen, 1983 ♀, ♂, USA | 2 | 2 | 9% | Sphaeropthalminae | Sphaeropthalmini |
Odontotilla Bischoff, 1920: *O. bidentata (André, 1905) ♀, ♂, South Africa | 1 | 1 | 0% | Myrmillinae | Myrmillinae |
Orientilla Lelej, 1979: O. aureorubra (Sichel et Radoszkowski, 1870) ♀, ♂, Sri Lanka; O. desponsa (Smith, 1855) ♀, ♂, Vietnam; O. kallata (Nurse, 1902) ♀, ♂, Sri Lanka; O. krombeini Lelej, 1998 ♀, ♂, Vietnam; O. sp. ♂, Vietnam; O. tausignata (Chen, 1957) ♀, China | 5 | 5 | 8% | Dasylabrini | Dasylabrinae |
Paramyrmosa de Saussure, 1880: P. brunnipes (Lepeletier, 1845) ♀, ♂, Austria, Serbia; P. pulla (Nylander, 1847) ♀, Russia | 2 | 1 | 1% | Myrmosini | Myrmosinae |
Pertyella Mickel, 1952: P. ? beata (Cameron, 1894) ♀, ♂, Panama; P. holmbergii (E.Lynch Arribálzaga, 1878) ♀, Argentina; P. ? salutatrix (Smith, 1879) ♀, ♂, Costa Rica; P. nr. lenti Casal, 1964 ♀, Argentina; P. sp. ♂, Peru | 4 | 3 | 2% | Pseudomethocina | Pseudomethocini |
Pherotilla Brothers, 2015: *P. mlanjeana (Bischoff, 1920) ♀, ♂, Malawi; P. oceanica (Mickel, 1935) ♀, ♂, Brunei; P. rufitincta (Hammer, 1957) ♀, ♂, Kenya | 3 | 3 | 10% | Rhopalomutillinae | |
Photomorphus Viereck, 1903: P. (P.) alogus Viereck, 1903 ♀, ♂, USA; P. (P.) myrmicoides (Cockerell, 1895) ♀, USA; P. (P.) quintilis (Viereck, 1906) ♂, USA | 2 | 2 | 0% | Sphaeropthalminae | Sphaeropthalmini |
Physetopoda Schuster, 1949: P. halensis (Fabricius, 1787) ♀, ♂, Kazakhstan; P. pierrei (Suárez, 1958) ♂, Mauritania, Chad; P. punctata (Latreille, 1792) ♀, ♂, Spain; P. portschinskii (Radoszkowski, 1888) ♂, Kazakhstan; P. scutellaris (Latreille, 1792) ♂, Kazakhstan; P. daghestanica (Radoszkowski, 1885) ♂, Kazakhstan, Ukraine | 2 | 6 | 7% | Smicromyrmina | Smicromyrmini |
Platymyrmilla André, 1900: *P. quinquefasciata (Olivier, 1811) ♀, ♂, Armenia, Ukraine | 1 | 1 | 0% | Myrmillinae | Myrmillinae |
Pristomutilla Ashmead, 1903: P. dentidorsis (André, 1908) ♀, Malawi; P. meigangana Nonveiller, 1995 ♂, Cameroon; P. nr. ctenophora Bischoff, 1921 ♂, South Africa; P. spp. ♀♀, ♂, South Africa, Tanzania | 3 | 3 | 7% | Smicromyrmina | Myrmillinae |
Promecilla André, 1902: P. decora (Smith, 1879), comb. n. ♀, ♂, Malaysia; *P. regia (Smith, 1855) ♀, ♂, India; P. spp. ♀♀, India | 4 | 2 | 8% | Smicromyrmina | Smicromyrmini |
Protophotopsis Schuster, 1947: *P. (P.) veneraria (Melander, 1903) ♀, ♂, USA | 1 | 1 | 0% | Sphaeropthalminae | Sphaeropthalmini |
Pseudocephalotilla Bischoff, 1920: P. atropos kalahariensis (Bischoff, 1921), comb. n. ♂, South Africa; P. beira (Péringuey, 1914), comb. n. ♀, ♂, South Africa; *P. beirana Bischoff, 1921, Mozambique; P. tettensis brunni (Bischoff, 1921), comb. n. ♀, South Africa | 2 | 3 | 7% | Smicromyrmina | Mutillini |
Pseudomethoca Ashmead, 1896: *P. frigida (Smith, 1855) ♀, ♂, USA; P. harpalyce (Fox, 1899) ♀, USA; P. oceola (Blake, 1871) ♂, USA; P. oculata (Banks, 1921) ♀, USA; P. propinqua (Cresson, 1865) ♀, ♂, USA; P. ravula (Cameron, 1894) ♀, Mexico; P. sanbornii (Blake, 1871) ♂, USA | 5 | 4 | 18% | Pseudomethocina | Pseudomethocini |
Pseudomutilla Costa, 1885: *Myrmilla (P.) capitata (Lucas, 1849) ♀, ♂, Italy, Spain; Myrmilla (P.) mavromoustakisi Hammer, 1950 ♀, ♂, Cyprus | 2 | 2 | 3% | Myrmillinae | Myrmillinae |
Pseudomyrmosa Suárez, 1980: P. gobicola Lelej, 1981, ♀, ♂, Russia; *P. minuta (Morawitz, 1894) ♀, ♂, Russia; P. schlettereri (Morawitz, 1890) ♀, ♂, Turkmenistan | 3 | 3 | 3% | Kudakrumiini | Kudakrumiinae |
Pseudophotopsis Andre, 1896: P. binghami Bischoff, 1920 ♂, United Arab Emirates; P. continua (Fabricius, 1804) ♀, ♂, Cameroon; *P. komarovii (Radoszkowski, 1885) ♀, ♂, Turkmenistan; P. schachruda (Skorikov, 1935) ♀, ♂, Cyprus; P. irana (Skorikov, 1935) ♂, Iran | 3 | 5 | 5% | Pseudophotopsidinae | Pseudophotopsidinae |
Radoszkowskitilla Lelej, 2005: *R. ceylonica (Lelej, 1993) ♀, India, Sri Lanka; R. karnataka Lelej, 2005 ♂, India; R. sinhala Lelej, 2005 ♂, Sri Lanka; R. tamila Lelej, 2005 ♂, Sri Lanka | 1 | 3 | 1% | Smicromyrmina 1 | Petersenidiini 1 |
Reedomutilla Mickel, 1964: R. dureti Casal, 1968 ♀, Argentina; R. fritzi Casal, 1968 ♂, Argentina; *R. gayi (Spinola) ♀, ♂, Chile; R. heraldica (Smith, 1855) ♀, Argentina; R. pubescens (Smith, 1875) ♂, Argentina | 3 | 3 | 6% | Sphaeropthalminae | Sphaeropthalmini |
Rhopalomutilla André, 1901: R. anguliceps (André, 1909) ♀, ♂, South Africa; R. carinaticeps Bischoff, 1920 ♀, ♂, Kenya, South Africa, Togo; *R. clavicornis (André, 1901) ♂, Zimbabwe | 2 | 3 | 7% | Rhopalomutillinae | |
Rimulotilla Brothers, 2015: R. conifera (Bischoff, 1920) ♀, ♂, Kenya; *R. tongaana (Péringuey, 1909) ♀, ♂, South Africa | 2 | 2 | 2% | Rhopalomutillinae | Rhopalomutillinae |
Ronisia Costa, 1858: *R. b. brutia (Petagna, 1787) ♀, ♂, Austria, Malta | 1 | 1 | 0% | Mutillina | Mutillini |
Scaptodactyla Burmeister, 1875: *S. ? heterogama Burmeister, 1875 ♀, ♂, Argentina | 1 | 1 | 0% | Sphaeropthalminae | Sphaeropthalmini |
Seyrigilla Krombein, 1972: *Stenomutilla (S.) nigroaurea (Sichel & Radoszkowski, 1869) ♂, Madagascar; Stenomutilla (S.) splendida Olsoufieff, 1938 ♀, Madagascar | 1 | 1 | 0% | Dasylabrini | Dasylabrinae |
Smicromyrme Thomson, 1870: S. bidenticulata Chen, 1957 ♂, Russia; S. lewisi Mickel, 1935 ♀, ♂, Russia, Japan; *S. rufipes (Fabricius, 1787) ♀, ♂, Austria, England | 2 | 3 | 3% | Smicromyrmina | Smicromyrmini |
Smicromyrmilla Suárez, 1965: S. ? alata (Bischoff, 1920) ♂, South Africa; S. tessmanni (Bischoff, 1920) ♀, Cameroon; S. spp. ♀♀♀♀, ♂, Lesotho, South Africa, Tanzania | 4 | 3 | 9% | Smicromyrmillini | Smicromyrmillini |
Sphaeropthalma Blake, 1871: Sphaeropthalma (S.) a. auripilis (Blake, 1871) ♀, ♂, USA; Sphaeropthalma (S.) pensylvanica floridensis Schuster, 1945 ♀, USA; Sphaeropthalma (S.) p. pensylvanica (Lepeletier, 1845) ♂, USA; *Sphaeropthalma (S.) pensylvanica scaeva (Blake,) ♂, USA | 2 | 3 | 2% | Sphaeropthalminae | Sphaeropthalmini |
Spinulomutilla Nonveiller, 1994: S. aureocincta (Magretti, 1884) ♀, ♂, Cameroon; S. braunsi (Bischoff, 1920) ♀, South Africa; *S. inaequalis Nonveiller, 1994 ♂, Cameroon; S. zoe (Péringuey, 1901) ♂, South Africa | 2 | 3 | 2% | Smicromyrmina 1 | Trogaspidiini 1 |
Stenomutilla André, 1896: *S. argentata (Villers, 1789) ♀, ♂, Italy, Spain; S. ? colligera (André, 1899) ♂, South Africa; S. eurydice (Péringuey, 1898) ♀, ♂, Namibia; S. hottentota (Fabricius, 1804) ♂, Malta; S. mlanjiana Bischoff, 1921 ♂, Zambia; S. nr. togoana Bischoff, 1921 ♀, Zambia; S. sp. ♂, Lesotho; S. tetrazonia Skorikov, 1935 ♀, ♂, Kazakhstan, Uzbekistan | 4 | 7 | 14% | Dasylabrini | Dasylabrinae |
Sulcotilla Bischoff, 1920: *S. sulcata (Magretti, 1884) ♀, ♂, Cameroon, Mali, Niger, Senegal | 1 | 1 | 0% | Smicromyrmina | Smicromyrmini |
Tallium André, 1902: T. catulus (Burmeister, 1875) ♀, ♂, Argentina; T. proseni Casal, 1965 ♀, Argentina; T. nr. precarium Suárez, 1960 ♂, Argentina; T. suarezi Casal, 1968 ♀, Argentina; T. tenebrosum (Gerstaecker, 1874) ♀, ♂, Argentina | 4 | 3 | 5% | Pseudomethocina | Pseudomethocini |
Timulla Ashmead, 1899: *T. dubitata (Smith, 1855) ♀, ♂, USA; T. ferrugata (Fabricius, 1804) ♀, ♂, USA; T. vagans (Fabricius, 1798) ♀, ♂, USA | 3 | 3 | 5% | Smicromyrmina 1 | Trogaspidiini 1 |
Tobantilla Casal, 1965: T. aleatrix Williams, Brothers& Pitts, 2011 ♀, Argentina; T. charrasca Casal, 1969 ♀, Argentina; T. drosa Williams, Brothers& Pitts, 2011 ♂, Argentina; T. ephemera Williams, Brothers& Pitts, 2011 ♂, Argentina; *T. montonera Casal, 1965 ♀, Argentina | 3 | 2 | 0% | Sphaeropthalminae | Sphaeropthalmini |
Tricholabiodes Radoszkowski, 1885: T. arabicus Suárez, 1967 ♂, United Arab Emirates; T. carinifer Bischoff, 1920 ♀, ♂, Namibia; T. ? lividus André, 1909 ♀, Namibia; T. semistriatus (Klug, 1829) ♀, Israel; T. nr. signatipennis (André, 1901) ♀, South Africa; T. sp. ♂, United Arab Emirates | 3 | 4 | 7% | Dasylabrini | Dasylabrinae |
Trispilotilla Bischoff, 1920: T. dewitziana (de Saussure, 1891) ♀, Mozambique; T. liopyga (Bischoff, 1920) ♀, South Africa; T. melanocephala Bischoff, 1920 ♂, Malawi; T. monteiroae Bischoff, 1920 ♂, South Africa; T. rugifera Nonveiller, 1973 ♀, Zimbabwe | 3 | 2 | 3% | Smicromyrmina 1 | Trogaspidiini 1 |
Trogaspidia Ashmead, 1899: T. fedtschenkoi (Radoszkowski, 1877) ♀, ♂, Turkmenistan, Uzbekistan; T. major Nonveiller & Petersen, 1995 ♀, ♂, South Africa; T. nr. caffrariae Bischoff, 1920 ♀, South Africa; T. themis (Péringuey, 1898) ♀, ♂, South Africa | 4 | 3 | 4% | Smicromyrmina 1 | Trogaspidiini 1 |
Tropidotilla Bischoff, 1920: T. cruenticeps (André, 1901) ♀, Cyprus; T. cypriadis Invrea, 1940 ♂, Cyprus; T. fimbriata (Klug, 1829) ♀, Eritrea; *T. litoralis (Petagna, 1787) ♀, ♂, Croatia, Greece, Spain; T. milmili (Magretti, 1898) ♂, Cameroon | 3 | 3 | 9% | Mutillina | Mutillini |
Vianatilla Casal, 1962: *V. nummularis (Gerstaecker, 1874) ♀, Argentina; V. spp., ♂♂, Costa Rica | 1 | 2 | 3% | Pseudomethocina | Pseudomethocini |
Viereckia Ashmead, 1903: V. ? acrisione (Péringuey, 1898) ♀, South Africa; V. ? nigra (Arnold, 1960) ♀, ♂, South Africa; V. spp. ♀♀♀, ♂♂♂, Lesotho, South Africa | 5 | 4 | 10% | Myrmillinae | Myrmillinae |
Wallacidia Lelej & Brothers, 2008: W. melmora (Cameron, 1905) ♂, Indonesia; W. philippinensis (Smith, 1855) ♀, ♂, Philippines; W. singapora (Mickel, 1935) ♂, Malaysia | 1 | 3 | 2% | Smicromyrmina 1 | Trogaspidiini 1 |
Xystromutilla André, 1905: *X. asperiventris André, 1905 ♀, ♂, Brazil; X. turrialba Casal, 1969 ♀, ♂, Panama | 2 | 2 | 11% | Sphaeropthalminae | Sphaeropthalmini |
Yamanetilla Lelej, 1996: Y. cassiope (Smith, 1879) ♂, Malaysia; Y. ? cassiope (Smith, 1879) ♀, Malaysia; *Y. nipponica (Tsuneki, 1972) ♀, ♂, Japan; Y. pedaria (Mickel, 1934) ♀, ♂, Philippines, Vietnam; Y. spp. ♀, ♂, Laos, Malaysia | 4 | 4 | 2% | Mutillina | Odontomutillini |
262 | 269 |
Characters and states for phylogenetic analysis of sub/genera of Mutillidae and four outgroup taxa
All characters are additive/ordered, unless otherwise stated; characters optimized as “fast”/“accelerated” (favouring reversals), except for those considered unlikely to show reversals, and therefore optimized as “slow”/“delayed” (favouring convergences) in Figs
1. Both sexes — Eye, pubescence and pores: 0 = Both present; 1 = Pubescence absent, pores present; 2 = Both absent. [0%; length = 6, ci = 0.33, ri = 0.80]
2. Both sexes — Pronotum, latero-ventral pubescent pit: 0 = Absent; 1 = Present. [0%; length = 1, uninformative]
3. Both sexes — Metasternum, level: 0 = Not depressed; 1 = Depressed. [0%; length = 1, uninformative]
4. Both sexes — Metasternum, form: 0 = Simple and flattened; 1 = With Y- to V-shaped carina or ridge, posterior arms leading to metacoxae bounding posterior median depression; 2 = With posterior median process(es) only. (NONADDITIVE) [1%; length = 1, ci = 1.00, ri = 1.00]
5. Both sexes — Metacoxa, postero-dorsally: 0 = Simple; 1 = With carinate tubercle; 2 = With lamellate process. [2%; length = 3, ci = 0.66, ri = 0.94]
6. Both sexes — Sternum I, posterolateral rounded densely pubescent depression: 0 = Absent; 1 = Present. [0%; length = 1, uninformative]
7. Both sexes — Tergum II and sternum I: 0 = Not articulated; 1 = Articulated, tergum II overlying lateral extremities of sternum I. [0%; length = 1, ci = 1.00, ri = 1.00]
8. Female — Head, shape: 0 = Normal, transverse, rounded posterolaterally; 1 = Normal, long, rounded posterolaterally; 2 = Broad, transverse, rounded posterolaterally; 3 = Broad, long, rounded posterolaterally; 4 = Broad, long, rectangular posterolaterally; 5 = Broad, transverse, protuberant/angular posterolaterally. (NONADDITIVE) [7%; length = 26, ci = 0.19, ri = 0.46]
9. Female — Occipital carina: 0 = Distinct and reflexed, complete; 1 = Distinct and reflexed, dorsal only; 2 = Absent, or not reflexed and scarcely discernible. [4%; length = 25, ci = 0.08, ri = 0.45]
10. Female — Eye, form: 0 = Weakly convex, following head contour; 1 = Moderately convex, distinct from head contour; 2 = Strongly convex, disjunct from head contour. [4%; length = 18, ci = 0.11, ri = 0.66]
11. Female — Eye, shape: 0 = Oval, inner margin more-or-less convex, long axis vertical; 1 = Oval, inner margin obviously sinuate or emarginate; 2 = Subcircular, inner margin convex, long axis vertical; 3 = Subcircular, inner margin convex, long axis horizontal. (NONADDITIVE) [10%; length = 13, ci = 0.23, ri = 0.82]
12. Female — Eye, surface: 0 = Ommatidia distinct; 1 = Ommatidia faintly distinguishable; 2 = Smooth, ommatidia not distinguishable. [3%; length = 13, ci = 0.15, ri = 0.63]
13. Female — Ocelli: 0 = Present, functional; 1 = Present but rudimentary; 2 = Absent. [1%; length = 4, ci = 0.50, ri = 0.80]
14. Female — Antennal socket, rim: 0 = Simple; 1 = Dorsally expanded as lamellate “tubercle” overhanging antennal base; 2 = Frons expanded as a ledge overhanging antennal socket. (NONADDITIVE) [0%; length = 3, ci = 0.66, ri = 0.50]
15. Female — Scape, radicle: 0 = Simple annular differentiation, not angled; 1 = Simple annular differentiation, angled; 2 = Flangelike expansion above radicle, angled. [0%; length = 2, ci = 1.00, ri = 1.00]
16. Female — Pedicel, length: 0 = Very short, <0.4 × length of flagellomere I; 1 = Short, >0.4 <0.7 × length of flagellomere I; 2 = About as long as flagellomere I. [4%; length = 23, ci = 0.08, ri = 0.51]
17. Female — Pedicel, shape: 0 = Shorter than wide; 1 = As long as wide; 2 = Longer than wide. [6%; length = 32, ci = 0.06, ri = 0.26]
18. Female — Flagellomere number: 0 = 10; 1 = 11. [0%; length = 1, ci = 1.00, ri = 1.00]
19. Female — Flagellomere I, length: 0 = Shorter than flagellomere II; 1 = 1–1.5 × length of flagellomere II; 2 = >1.8 × length of flagellomere II. [4%; length = 13, ci = 0.15, ri = 0.63]
20. Female — Flagellomere I, shape: 0 = Shorter than wide; 1 = About as long as wide; 2 = >1.3 <2.0 × as long as wide; 3 = >2 × as long as wide. [7%; length = 25, ci = 0.12, ri = 0.48]
21. Female — Genal carina: 0 = Absent; 1 = Present but weak; 2 = Present and strong. [11%; length = 28, ci = 0.07, ri = 0.54]
22. Female — Genal carina, extent (carina absent = -): 0 = Ending distant from hypostomal carina; 1 = Ending adjacent to hypostomal carina; 2 = Continuous with hypostomal carina. [3%; length = 17, ci = 0.11, ri = 0.37]
23. Female — Genal carina, armature (carina absent = -): 0 = Carina simple, unarmed; 1 = With small lamellate tooth anteroventrally; 2 = With strong conical or pyramidal tooth anteroventrally; 3 = With teeth posterodorsally and anteroventrally. (NONADDITIVE) [3%; length = 6, ci = 0.50, ri = 0.40]
24. Female — Hypostomal carina: 0 = Complete, simple; 1 = Complete, flange-like; 2 = Complete, with distinct tooth laterally; 3 = Complete, with tooth/tubercle/elevation at about midlength; 4 = Strong anterolaterally but obsolete posteriorly; 5 = Strong posteriorly but absent anterolaterally. (NONADDITIVE) [7%; length = 25, ci = 0.20, ri = 0.53]
25. Female — Pleurostomal carina: 0 = Absent; 1 = Slight, ending at inner mandibular edge; 2 = Distinct, together with hypostomal carina forming curved to angulate ridge ending at outer mandibular articulation; 3 = Distinct, together with hypostomal carina forming straight ridge ending at outer mandibular articulation. [4%; length = 28, ci = 0.10, ri = 0.46]
26. Female — Postgenal carina/ridge: 0 = Absent; 1 = Distinct, separate from hypostomal carina; 2 = Distinct, merging with hypostomal carina. [4%; length = 15, ci = 0.13, ri = 0.66]
27. Female — Postmandibular carina (posteroventral to mandible base): 0 = Absent; 1 = Present, simple blunt ridge. [0%; length = 4, ci = 0.25, ri = 0.50]
28. Female — Oral and mandibular fossae: 0 = Continuous, junction about half mandible width or more; 1 = Continuous, junction much narrowed; 2 = Separated by anteriorly unfused depressed cuticular bridge; 3 = Separated by anteriorly fused much-depressed cuticular bridge; 4 = Separated by anteriorly fused superficial cuticular bridge. [3%; length = 26, ci = 0.15, ri = 0.78]
29. Female — Mandible, apical teeth: 0 = Three; 1 = Two; 2 = One. [7%; length = 23, ci = 0.08, ri = 0.40]
30. Female — Mandible, shape: 0 = Apically not expanded; 1 = Apically expanded. [3%; length = 7, ci = 0.14, ri = 0.33]
31. Female — Mandible, posteroventral basal expansion: 0 = Absent; 1 = Present, toothlike; 2 = Present, flangelike, apically abrupt; 3 = Present, flangelike, apically oblique. (NONADDITIVE) [5%; length = 11, ci = 0.27, ri = 0.00]
32. Female — Mandible, inner basal tooth: 0 = Absent; 1 = Present, acute; 2 = Present, mediobasal obtuse flange. (NONADDITIVE) [2%; length = 13, ci = 0.15, ri = 0.35]
33. Female — Labio-maxillary complex: 0 = Short; 1 = Elongated prementum and stipes. [0%; length = 1, ci = 1.00, ri = 1.00]
34. Female — Prementum: 0 = Evenly convex to weakly medio-longitudinally carinate; 1 = With posteromedian domelike tubercle or elevation; 2 = With sharp posteromedian elevation; 3 = With anteriorly indented posteromedian domelike elevation; 4 = Flattened, depressed to weakly concave; 5 = Longitudinally convex with deep narrow anteromedian groove; 6 = With strong long narrow median carina; 7 = With paired medial longitudinal carinae. (NONADDITIVE) [3%; length = 14, ci = 0.50, ri = 0.50]
35. Female — Maxillary palp, segments: 0 = Six-segmented; 1 = Two-segmented; 2 = Unsegmented. [2%; length = 2, ci = 1.00, ri = 1.00]
36. Female — Maxillary palp, length: 0 = Shorter than fore tibia; 1 = 1–1.5 × length fore tibia; 2 = >1.5 <2 × length fore tibia; 3 = >2 × length fore tibia. [10%; length = 28, ci = 0.10, ri = 0.37]
37. Female — Labial palp, segments: 0 = Four-segmented; 1 = Two-segmented; 2 = Unsegmented. [2%; length = 2, ci = 1.00, ri = 1.00]
38. Female — Wings: 0 = Present; 1 = Absent. [0%; length = 1, ci = 1.00, ri = 1.00]
39. Female — Mesosomal dorsum, flattened decumbent setae: 0 = Absent; 1 = Present, laterally flattened, slender, arcuate; 2 = Present, laterally flattened, broad, lanceolate; 3 = Present, dorsoventrally flattened, slender, arcuate. (NONADDITIVE) [0%; length = 8, ci = 0.37, ri = 0.44]
40. Female — Mesosoma, form (dorsal view; winged = -): 0 = More or less parallel-sided; 1 = Mesothorax protuberant well anterior to metathoracic spiracle, propodeum narrower than prothorax; 2 = Mesothorax protuberant just anterior to metathoracic spiracle, propodeum narrower than prothorax; 3 = Ovate, propodeum about as broad as prothorax; 4 = Mesothorax margin straightish, propodeum much broader than prothorax; 5 = Mesothorax margin dorsally concave, pronotum broadest; 6 = Pronotum broadest, mesothoracic margin straightish, mesosoma evenly narrowed posteriorly. (NONADDITIVE) [6%; length = 15, ci = 0.40, ri = 0.81]
41. Female — Mesosoma, dorsolateral margin: 0 = Smooth, sinuate or weakly tuberculate; 1 = With distinct teeth. [5%; length = 11, ci = 0.09, ri = 0.33]
42. Female — Pro-mesonotal suture: 0 = Distinct and freely articulating; 1 = Distinct but fused, not articulating; 2 = Obliterated or very indistinct and fused, not articulating. [1%; length = 2, ci = 1.00, ri = 1.00]
43. Female — Pronotum, lateral length: 0 = About as long as distance between pronotal and propodeal spiracles; 1 = <0.8 × distance between pronotal and propodeal spiracles. [2%; length = 8, ci = 0.12, ri = 0.61]
44. Female — Pronotum, humeral angle: 0 = Rounded; 1 = Abrupt; 2 = Vertically carinate to weakly dentate; 3 = With prominent tooth or spine. [8%; length = 43, ci = 0.06, ri = 0.33]
45. Female — Pronotum, dorsolateral setose area/epaulet: 0 = Absent; 1 = Present, dispersed patch; 2 = Present, clearly delimited tubercle/tuft. [3%; length = 21, ci = 0.09, ri = 0.71]
46. Female — Pronotum, anterodorsal setose area/epaulet: 0 = Absent; 1 = Present, dispersed patch; 2 = Present, clearly delimited tubercle/tuft. [5%; length = 22, ci = 0.09, ri = 0.67]
47. Female — Pronotum, posteroventral margin: 0 = Distinct and complete; 1 = Indistinct or interrupted; 2 = Obliterated. [4%; length = 27, ci = 0.07, ri = 0.59]
48. Female — Meso-metanotal suture: 0 = Distinct; 1 = Obliterated or very indistinct. [0%; length = 2, ci = 0.50, ri = 0.75]
49. Female — Mesosoma, scutellar scale (winged = -): 0 = Absent; 1 = Present. [2%; length = 11, ci = 0.09, ri = 0.61]
50. Female — Propodeum, shape: 0 = >0.6 × as long as wide; 1 = <0.6 × as long as wide. [4%; length = 26, ci = 0.03, ri = 0.35]
51. Female — Propodeum, posterodorsal margin, form: 0 = Smoothly rounded; 1 = Abrupt but not ridgelike; 2 = Carinate or ridgelike. [3%; length = 28, ci = 0.07, ri = 0.51]
52. Female — Propodeum, posterodorsal margin, dentition: 0 = Smooth or tuberculate; 1 = With one weak median spine or vertical tubercle; 2 = With two lateral spines or teeth only; 3 = With three spines; 4 = With more than three spines; 5 = With two median teeth and two lateral spines or teeth; 6 = With two large sublateral cylindrical spines. (NONADDITIVE) [2%; length = 14, ci = 0.42, ri = 0.38]
53. Female — Propodeum, posterolateral margin: 0 = Smooth or tuberculate; 1 = Dentate or spinose. [0%; length = 10, ci = 0.10, ri = 0.25]
54. Female — Mesopleuron, dorsal region: 0 = Strongly protuberant; 1 = Weakly convex; 2 = Depressed. [4%; length = 22, ci = 0.09, ri = 0.73]
55. Female — Mesopleural ridge (usually setose): 0 = Absent; 1 = Indistinct and joined to mesonotal tubercle; 2 = Strong and joined to mesonotal tubercle; 3 = Joined to pronotal spiracle; 4 = Present only ventrally, with a narrow dorsal ridge to mesonotal tubercle; 5 = Present only ventrally, with a narrow dorsal ridge to pronotal spiracle; 6 = Present only ventrally; 7 = Ventrally evanescent, a dorsal ridge to pronotal spiracle; 8 = Entirely indistinct, joined to pronotal spiracle; 9 = A fine ridge approaching pronotal spiracle. (NONADDITIVE) [10%; length = 34, ci = 0.26, ri = 0.62]
56. Female — Mesopleural ridge, ventral section, position (absent = -): 0 = Anterior to midpoint of mesocoxa; 1 = Dorsal to midpoint of mesocoxa. [1%; length = 2, ci = 0.50, ri = 0.66]
57. Female — Mesopleural ridge, ventral section, form (absent = -): 0 = Blunt; 1 = Sharply carinate. [4%; length = 9, ci = 0.11, ri = 0.68]
58. Female — Meso-metapleural suture, direction (indistinguishable = -): 0 = Weakly curved (separate from mesopleural ridge); 1 = Strongly angled (joining mesopleural ridge). [2%; length = 4, ci = 0.25, ri = 0.91]
59. Female — Meso-metapleural suture, development: 0 = Distinct; 1 = Distinct ventrally only; 2 = Obliterated on surface. [4%; length = 18, ci = 0.11, ri = 0.38]
60. Female — Meso-metapleural “bridge”: 0 = Absent; 1 = Present. [0%; length = 1, ci = 1.00, ri = 1.00]
61. Female — Metapleural-propodeal suture, development: 0 = Entirely distinct; 1 = Obliterated dorsally, distinct ventral to endophragmal pit; 2 = Obliterated dorsally, vague ventral to endophragmal pit; 3 = Entirely obliterated on surface; 4 = Distinct dorsally, obliterated ventral to endophragmal pit. (NONADDITIVE) [11%; length = 29, ci = 0.13, ri = 0.55]
62. Female — Mesosternum just anterior to mesocoxae: 0 = Smoothly rounded; 1 = With paired transverse/oblique carinae (may be toothed mesally); 2 = With paired lamellate projections mesally. (NONADDITIVE) [1%; length = 2, uninformative]
63. Female — Mesocoxae, contiguity: 0 = Contiguous mesally; 1 = Slightly separated mesally. [1%; length = 3, ci = 0.33, ri = 0.71]
64. Female — Metasternum, posterior median process (absent = -): 0 = Shorter than coxal height, tridentate; 1 = Shorter than coxal height, shallowly bidentate; 2 = Shorter than coxal height, deeply bidentate; 3 = Shorter than coxal height, unidentate; 4 = Longer than coxal height, tridentate; 5 = Longer than coxal height, acutely unidentate; 6 = Longer than coxal height, obtusely unidentate; 7 = Shorter than coxal height, a transverse crenulate ridge. (NONADDITIVE) [5%; length = 28, ci = 0.25, ri = 0.58]
65. Female — Metacoxal cavities: 0 = Open; 1 = Partially closed; 2 = Closed. [0%; length = 3, ci = 0.66, ri = 0.80]
66. Female — Tarsomeres, apicoventral median ovoid pulvillus: 0 = Absent; 1 = On tarsomeres I–IV. [0%; length = 1, uninformative]
67. Female — Tarsal claws: 0 = Midventrally toothed; 1 = Simple; 2 = Apically deeply bifid. (NONADDITIVE) [0%; length = 3, ci = 0.66, ri = 0.66]
68. Female — Arolia: 0 = Present; 1 = Absent. [0%; length = 2, ci = 0.50, ri = 0.80]
69. Female — Fore tibia, inner (anterior) secretory structure: 0 = None; 1 = Broad coarsely setose delimited patch; 2 = Linear to oval finely perforated depression; 3 = Vertically elongate groove/pore; 4 = Obliquely elongate groove/pore; 5 = Obliquely oval to circular pore; 6 = Obliquely elongate groove/pore and linear finely perforated depression; 7 = Two apical separated obliquely oval pores; 8 = Linear to oval finely perforated convexity; 9 = Basal elongate/oval and separated apical round pores. (NONADDITIVE) [6%; length = 27, ci = 0.33, ri = 0.64]
70. Female — Fore tibia, outer (posterior) secretory structure: 0 = None; 1 = Linear to oval finely perforated depression; 2 = Vertically elongate groove/pore; 3 = Obliquely elongate groove/pore; 4 = Obliquely oval to circular pore. (NONADDITIVE) [3%; length = 17, ci = 0.17, ri = 0.36]
71. Female — Fore calcar blade: 0 = Linearly narrow, margin entire; 1 = Linearly narrow, margin finely pectinate; 2 = Expanded, longish >0.4 × length of calcar; 3 = Expanded, almost square, <0.4 × length of calcar; 4 = Concave, narrow, apically expanded. (NONADDITIVE) [0%; length = 6, ci = 0.66, ri = 0.84]
72. Female — Meso- and metatibial articulated spines, mean number: 0 = 0–4; 1 = 5–9; 2 = 10–14; 3 = >14. [12%; length = 28, ci = 0.10, ri = 0.40]
73. Female — Metacoxa, mesally: 0 = Smoothly rounded; 1 = Longitudinally carinate. [5%; length = 9, ci = 0.11, ri = 0.80]
74. Female — Metatibia, apex dorsally: 0 = Evenly rounded; 1 = With elevated tubercle bearing spine(s); 2 = With cylindrical process bearing spine. [3%; length = 9, ci = 0.22, ri = 0.22]
75. Female — Metatibia, posterior (inner) surface: 0 = Convex, setose, punctate; 1 = Flattened and broadened, with smooth delimited area. [0%; length = 1, ci = 1.00, ri = 1.00]
76. Female — Metatibia, posteroapical secretory structure: 0 = Absent; 1 = Present, delimited patch of dense setae; 2 = Present, linear setose felt-line-like; 3 = Present, a small pore; 4 = Present, a deep narrow longitudinal groove. (NONADDITIVE) [3%; length = 12, ci = 0.33, ri = 0.63]
77. Female — Metatibia, apical spurs: 0 = Both similar, unmodified; 1 = Inner modified as cleaner. [0%; length = 2, ci = 0.50, ri = 0.80]
78. Female — Tergum I, profile: 0 = Broadly convex; 1 = Anterior and dorsal faces merging; 2 = Anterior and dorsal faces distinct, bounded. [10%; length = 30, ci = 0.06, ri = 0.37]
79. Female — Tergum I, base: 0 = Simple; 1 = With paired vertical ridges; 2 = With paired expanded “auricles”. (NONADDITIVE) [2%; length = 3, ci = 0.66, ri = 0.88]
80. Female — Tergum I, shape: 0 = ≥0.5 × length of tergum II, gradually broadened posteriorly, sessile on tergum II; 1 = ≥0.4 × length of tergum II, strongly broadened, parallel-sided posteriorly, discontinuous with tergum II; 2 = <0.5 × length of tergum II, gradually broadened posteriorly, sessile on tergum II; 3 = <0.5 × length of tergum II, gradually broadened posteriorly, constricted apically, disjunct from tergum II; 4 = <0.25 × length of tergum II, entirely parallel-sided, discontinuous with tergum II. (NONADDITIVE) [4%; length = 11, ci = 0.36, ri = 0.63]
81. Female — Tergum I, apical width: 0 = >0.75 × width of tergum II; 1 = <0.75 >0.5 × width of tergum II; 2 = <0.5 × width of tergum II. [5%; length = 21, ci = 0.09, ri = 0.51]
82. Female — Tergum I and propodeum, pubescence: 0 = All simple; 1 = Some brachyplumose; 2 = Some fully plumose. [2%; length = 6, ci = 0.33, ri = 0.87]
83. Female — Tergum I apex, pale pubescent markings: 0 = None; 1 = Median pale spot; 2 = Paired pale spots; 3 = Pale band. (NONADDITIVE) [15%; length = 31, ci = 0.09, ri = 0.39]
84. Female — Tergum II, length: 0 = <0.75 × length of terga III–VI; 1 = 0.75–1.25 × length of terga III–VI; 2 = >1.25 × length of terga III–VI. [1%; length = 10, ci = 0.20, ri = 0.73]
85. Female — Tergum II, pale markings, number: 0 = None; 1 = Odd number (unpaired); 2 = Even number (paired); 3 = Broad band. (NONADDITIVE) [13%; length = 21, ci = 0.14, ri = 0.63]
86. Female — Tergum II, pale markings, composition (absent = -): 0 = Pubescence only; 1 = Integumental. [6%; length = 7, ci = 0.14, ri = 0.50]
87. Female — Tergum II, apical fringe setae: 0 = Entirely simple; 1 = Some densely plumose. [0%; length = 1, ci = 1.00, ri = 1.00]
88. Female — Tergum II, felt line, presence: 0 = Absent; 1 = Present = 1. [1%; length = 4, ci = 0.25, ri = 0.81]
89. Female — Tergum II, felt line, form (absent = -): 0 = Linear and superficial; 1 = Broad lateral patch; 2 = Invaginated (elongate or pitlike); 3 = Small indefinite anterior patch. (NONADDITIVE) [0%; length = 3, ci = 1.00, ri = 1.00]
90. Female — Tergum III, stridulitrum: 0 = Absent; 1 = Present. [0%; length = 4, ci = 0.25, ri = 0.66]
91. Female — Tergum VI, form: 0 = Entirely evenly sculptured; 1 = Evenly sculptured except apical area much finer/smoother; 2 = With smooth(ish) unbounded longitudinal median area, laterally sculptured; 3 = With distinct bounded pygidial plate. (NONADDITIVE) [6%; length = 19, ci = 0.15, ri = 0.52]
92. Female — Sternum I, differentiation: 0 = Smoothly overlapping sternum II; 1 = Briefly declivous and abutting sternum II; 2 = Depressed posteriorly, constricted and abutting sternum II. [1%; length = 2, ci = 1.00, ri = 1.00]
93. Female — Sternum II, felt line, presence: 0 = Absent; 1 = Present. [3%; length = 5, ci = 0.20, ri = 0.42]
94. Female — Sternum II, felt line, form (absent = -): 0 = Dispersed traces only; 1 = Distinctly compact and linear. [1%; length = 1, uninformative]
95. Female — Sternum VI, sting aperture: 0 = Lateral areas differentiated, sting aperture slit-like; 1 = Lateral areas dorsomesally produced, sting aperture circular. [0%; length = 1, ci = 1.00, ri = 1.00]
96. Female — Sternum VI, armature: 0 = Without processes; 1 = With pair of apical processes (apex notched); 2 = With pair of acute lateral teeth basally; 3 = With two pairs of lateroventral tubercles; 4 = With two pairs of apical processes/teeth (apex 4-lobed). (NONADDITIVE) [8%; length = 22, ci = 0.18, ri = 0.41]
97. Female — Gonapophysis IX, sting curvature: 0 = Weakly convexly arcuate dorsally; 1 = Strongly convexly arcuate dorsally, apex directed downwards. [0%; length = 2, ci = 0.50, ri = 0.50]
98. Male — Head, width across mandibular bases: 0 = <0.6 × maximum head width; 1 = >0.6 × maximum head width. [1%; length = 4, ci = 0.25, ri = 0.57]
99. Male — Head, pubescence: 0 = Entirely simple; 1 = Some brachyplumose; 2 = Some fully plumose. [2%; length = 6, ci = 0.33, ri = 0.88]
100. Male — Occipital carina: 0 = Distinct and reflexed, complete; 1 = Distinct and reflexed, dorsal only; 2 = Absent, or not reflexed and scarcely discernible. [5%; length = 17, ci = 0.11, ri = 0.28]
101. Male — Eye, form: 0 = Weakly convex, following head contour; 1 = Moderately convex, distinct from head contour; 2 = Strongly convex, disjunct from head contour. [5%; length = 21, ci = 0.09, ri = 0.66]
102. Male — Eye, shape: 0 = Broadly oval, inner margin convex to weakly sinuate; 1 = Broadly oval, inner margin acutely but shallowly emarginate; 2 = Broadly oval, inner margin acutely and deeply emarginate; 3 = Subcircular, inner margin sinuate to weakly emarginate; 4 = Subcircular, inner margin acutely and deeply emarginate; 5 = Subcircular, inner margin roughly convex, long axis vertical; 6 = Subcircular, inner margin roughly convex, long axis horizontal. (NONADDITIVE) [6%; length = 19, ci = 0.31, ri = 0.82]
103. Male — Eye, surface: 0 = Ommatidia distinct; 1 = Ommatidia faintly distinguishable; 2 = Smooth, ommatidia not distinguishable. [2%; length = 13, ci = 0.15, ri = 0.31]
104. Male — Ocelli: 0 = Present, normal; 1 = Present, much enlarged; 2 = Absent. (NONADDITIVE) [3%; length = 6, ci = 0.33, ri = 0.20]
105. Male — Antennal socket, rim: 0 = Simple; 1 = Dorsally expanded as lamellate “tubercle” overhanging antennal base; 2 = Frons expanded as a ledge overhanging antennal socket. (NONADDITIVE) [0%; length = 2, ci = 1.00, ri = 1.00]
106. Male — Antennal scrobe, dorsal secretory tubercle and carina: 0 = No tubercle, no carina; 1 = Weak transverse carina only; 2 = Strong transverse carina only; 3 = Secretory tubercle only; 4 = Secretory tubercle and weak transverse carina; 5 = Secretory tubercle and strong transverse carina. (NONADDITIVE) [19%; length = 37, ci = 0.13, ri = 0.54]
107. Male — Scape, radicle: 0 = Simple annular differentiation, not angled; 1 = Simple annular differentiation, angled; 2 = Flangelike expansion above radicle, angled. [0%; length = 3, ci = 0.66, ri = 0.83]
108. Male — Scape, ventral ridges: 0 = None; 1 = One only; 2 = Two, one less developed; 3 = Two, equally well developed. [11%; length = 38, ci = 0.07, ri = 0.58]
109. Male — Pedicel, length: 0 = Very short, <0.4 × length of flagellomere I; 1 = Short, >0.4 <0.8 × length of flagellomere I; 2 = About as long as flagellomere I. [10%; length = 24, ci = 0.08, ri = 0.35]
110. Male — Pedicel, shape: 0 = Distinctly shorter than wide; 1 = About as long as wide; 2 = Distinctly longer than wide. [12%; length = 25, ci = 0.08, ri = 0.37]
111. Male — Flagellomere I, length: 0 = <0.6 × length of flagellomere II; 1 = Subequal to flagellomere II; 2 = >1.3 × length of flagellomere II. [6%; length = 15, ci = 0.13, ri = 0.43]
112. Male — Flagellomere I, shape: 0 = Wider than long; 1 = About as long as wide; 2 = >1.3 <2.0 × as long as wide; 3 = >2 × as long as wide. [12%; length = 30, ci = 0.10, ri = 0.54]
113. Male — Flagellomere I, form: 0 = Cylindrical; 1 = Weakly flattened ventrally only; 2 = Strongly flattened and broadened. [1%; length = 4, ci = 0.50, ri = 0.85]
114. Male — Head, genal carina: 0 = Absent; 1 = Present but weak; 2 = Present and strong. [10%; length = 12, ci = 0.16, ri = 0.16]
115. Male — Head, genal carina, extent (carina absent = -): 0 = Ending distant from hypostomal carina; 1 = Ending adjacent to or continuous with hypostomal carina. [1%; length = 1, uninformative]
116. Male — Head, genal carina, armature (carina absent = -): 0 = Carina simple, unarmed; 1 = With small lamellate tooth anteroventrally; 2 = With strong conical tooth anteroventrally; 3 = With teeth posterodorsally and anteroventrally; 4 = With strong short lamellate tooth posterodorsally. (NONADDITIVE) [3%; length = 3, ci = 0.66, ri = 0.50]
117. Male — Hypostomal carina: 0 = Complete, simple; 1 = Complete, flangelike; 2 = Complete, with tooth/tubercle/elevation at about midlength; 3 = Strong anterolaterally but obsolete posteriorly. (NONADDITIVE) [10%; length = 29, ci = 0.10, ri = 0.39]
118. Male — Pleurostomal carina: 0 = Absent; 1 = Slight, ending at inner mandibular edge; 2 = Distinct, together with hypostomal carina forming curved to angulate ridge ending at outer mandibular articulation; 3 = Distinct, together with hypostomal carina forming straight ridge ending at outer mandibular articulation. [4%; length = 26, ci = 0.11, ri = 0.28]
119. Male — Postgenal carina/ridge: 0 = Absent; 1 = Distinct, separate from hypostomal carina; 2 = Distinct, merging with hypostomal carina. [2%; length = 17, ci = 0.11, ri = 0.54]
120. Male — Postmandibular carina (posteroventral to mandible base): 0 = Absent; 1 = Present, simple blunt ridge; 2 = Present, broad smooth tubercle. (NONADDITIVE) [0%; length = 4, ci = 0.50, ri = 0.60]
121. Male — Oral and mandibular fossae: 0 = Continuous, junction about half mandible width or more; 1 = Continuous, junction much narrowed; 2 = Separated by anteriorly unfused depressed cuticular bridge; 3 = Separated by anteriorly fused much-depressed cuticular bridge; 4 = Separated by anteriorly fused superficial cuticular bridge. [1%; length = 14, ci = 0.28, ri = 0.88]
122. Male — Mandible, apical teeth: 0 = Four; 1 = Three; 2 = Two. [9%; length = 20, ci = 0.10, ri = 0.58]
123. Male — Mandible, shape: 0 = Apically not expanded; 1 = Apically expanded. [8%; length = 19, ci = 0.05, ri = 0.33]
124. Male — Mandible, posteroventral basal tooth: 0 = Absent; 1 = Present, small, toothlike; 2 = Present, enlarged, toothlike; 3 = Present, lamellate, apically oblique; 4 = Present, lamellate, apically abrupt. (NONADDITIVE) [10%; length = 26, ci = 0.11, ri = 0.39]
125. Male — Mandible, inner basal tooth: 0 = Absent; 1 = Present. [5%; length = 6, ci = 0.16, ri = 0.37]
126. Male — Prementum: 0 = Evenly convex or weakly medio-longitudinally carinate; 1 = With posteromedian domelike tubercle or elevation; 2 = With sharp posterior median elevation; 3 = With anteriorly indented posterior domelike elevation; 4 = Flattened, depressed to weakly concave; 5 = Longitudinally convex with deep narrow anteromedian groove; 6 = With strong long narrow median carina; 7 = With paired longitudinal carinae; 8 = Flattened with posterior median transversely lamellate projection. (NONADDITIVE) [5%; length = 19, ci = 0.36, ri = 0.36]
127. Male — Maxillary palp, length: 0 = Shorter than fore tibia; 1 = >1 <1.5 × length of fore tibia; 2 = >1.5 <2 × length of fore tibia; 3 = >2 × length of fore tibia. [5%; length = 28, ci = 0.10, ri = 0.43]
128. Male — Labial palp, mid segments: 0 = More or less cylindrical; 1 = Flattened and expanded, asymmetrical. [0%; length = 4, ci = 0.25, ri = 0.57]
129. Male — Mesosomal dorsum, decumbent setae: 0 = Cylindrical, slender, straight to weakly arcuate; 1 = Laterally flattened, slender, arcuate; 2 = Laterally flattened, broad, lanceolate. [10%; length = 12, ci = 0.16, ri = 0.61]
130. Male — Pro-mesonotal suture (indistinguishable = -): 0 = Weakly concave; 1 = Strongly but evenly concave; 2 = Broadly V-shaped (laterally straight, mesal third curved); 3 = Abruptly V-shaped (laterally straight, mesally angled). [10%; length = 29, ci = 0.10, ri = 0.61]
131. Male — Pronotum, humeral angle: 0 = Smoothly rounded; 1 = Blunt; 2 = Vertically carinate or dentate; 3 = Prominent tooth or spine. [10%; length = 36, ci = 0.08, ri = 0.34]
132. Male — Pronotum, dorsolateral setose area/epaulet: 0 = Absent; 1 = Present, dispersed patch; 2 = Present, clearly delimited tubercle/tuft. [6%; length = 12, ci = 0.16, ri = 0.75]
133. Male — Pronotum, anterodorsal setose area/epaulet: 0 = Absent; 1 = Present, dispersed patch; 2 = Present, clearly delimited tubercle/tuft. [9%; length = 16, ci = 0.12, ri = 0.67]
134. Male — Mesoscutum, length (apterous/brachypterous = -): 0 = Short anterior to tegulae; 1 = Extended far anterior to tegulae. [0%; length = 9, ci = 0.11, ri = 0.57]
135. Male — Mesoscutum, notaulus (apterous/brachypterous = -): 0 = Present and complete; 1 = Present but incomplete; 2 = Absent. [11%; length = 24, ci = 0.08, ri = 0.55]
136. Male — Mesoscutum, parapsidal furrow (apterous/brachypterous = -): 0 = Obvious, complete; 1 = Obvious posteriorly only, absent or a mere scar anteriorly; 2 = An obvious groove anteriorly only, absent posteriorly; 3 = Much reduced, at most a superficial scar anteriorly; 4 = Interrupted, a faint groove posteriorly, a superficial scar anteriorly. (NONADDITIVE) [8%; length = 21, ci = 0.19, ri = 0.62]
137. Male — Mesoscutum, posterolateral margin (apterous/brachypterous = -): 0 = Rounded; 1 = Lobed but flattish; 2 = Dentate and protruding upwards. [8%; length = 13, ci = 0.15, ri = 0.62]
138. Male — Mesoscutellum (apterous/brachypterous = -): 0 = Simple, even with metanotum; 1 = Pulvinate with smooth median ridge; 2 = Posteromesally produced (conical) with smooth median ridge; 3 = Laterally produced as curved posteriorly dentate flange; 4 = Posteriorly produced and overhanging metanotum; 5 = Swollen, discontinuous with metanotum. (NONADDITIVE) [5%; length = 17, ci = 0.29, ri = 0.47]
139. Male — Axilla, posterolateral dorsal margin (apterous/brachypterous = -): 0 = Rounded, posterolateral surface convex or flat; 1 = Narrowly rounded, posterolateral surface concave; 2 = Carinate, posterolateral surface concave; 3 = Flangelike, apex broadly obtuse; 4 = Flangelike, apex strongly dentate. [13%; length = 21, ci = 0.19, ri = 0.72]
140. Male — Axilla, anterolateral dorsal extremity (apterous/brachypterous = -): 0 = Evenly rounded (vertical ridge may be present ventrally); 1 = With slight vertical ridge broadening ventrally; 2 = With strong evenly developed vertical ridge or flange; 3 = With blunt tubercle or tooth dorsally; 4 = With abrupt broad vertical flange dorsally. [4%; length = 27, ci = 0.14, ri = 0.62]
141. Male — Tegula, length (apterous/brachypterous = -): 0 = Short, round, posteriorly distant from trans-scutal articulation; 1 = Slightly elongate, posteriorly reaching trans-scutal articulation or slightly beyond; 2 = Elongate, posteriorly extending well beyond trans-scutal articulation. [2%; length = 10, ci = 0.20, ri = 0.84]
142. Male — Tegula, form (apterous/brachypterous = -): 0 = Evenly convex to flattish posteriorly; 1 = Posteriorly recurved; 2 = Longitudinally angulate basally. (NONADDITIVE) [2%; length = 6, ci = 0.33, ri = 0.86]
143. Male — Tegula, free posterior inner margin (apterous/brachypterous = -): 0 = More or less straight or weakly convex; 1 = Distinctly concave. [1%; length = 5, ci = 0.20, ri = 0.87]
144. Male — Propodeum, length (apterous/brachypterous = -): 0 = About as long as high; 1 = Much shorter than high. [2%; length = 14, ci = 0.07, ri = 0.23]
145. Male — Propodeum, disc sculpture (apterous/brachypterous = -): 0 = Evenly sculptured; 1 = Larger basal fields and smaller distal fields; 2 = Three large fields bordered by strong longitudinal carinae. (NONADDITIVE) [5%; length = 9, ci = 0.22, ri = 0.83]
146. Male — Propodeum, disc and declivity (apterous/brachypterous = -): 0 = Broadly rounded; 1 = Distinct but merging; 2 = Abruptly differentiated. [10%; length = 24, ci = 0.08, ri = 0.38]
147. Male — Propodeum, dorsolateral margin (apterous/brachypterous = -): 0 = Rounded; 1 = Carinate or distinctly angled. [7%; length = 11, ci = 0.09, ri = 0.54]
148. Male — Propodeum, posterolateral margin: 0 = Smooth or tuberculate; 1 = Dentate or spinose. [2%; length = 2, ci = 0.50, ri = 0.50]
149. Male — Prepectus: 0 = Articulating with mesepisternum; 1 = Fused to mesepisternum. [0%; length = 1, uninformative]
150. Male — Meso-metapleural suture, fusion (apterous/brachypterous = -): 0 = Articulating; 1 = Immovable although not fused; 2 = Partially or entirely fused. [0%; length = 3, ci = 0.66, ri = 0.92]
151. Male — Meso-metapleural suture, shape (apterous/brachypterous = -): 0 = Entirely almost straight; 1 = Posteriorly convex; 2 = Sinuate, ventral section scarcely to strongly concave. (NONADDITIVE) [0%; length = 2, ci = 1.00, ri = 1.00]
152. Male — Meso-metapleural “bridge”: 0 = Absent; 1 = Present. [0%; length = 1, ci = 1.00, ri = 1.00]
153. Male — Metapleural-propodeal suture (apterous/brachypterous = -): 0 = Entirely distinct; 1 = Obliterated dorsal to endophragmal pit, distinct ventrally; 2 = Obliterated dorsal to endophragmal pit, vague ventrally; 3 = Entirely obliterated on surface. [20%; length = 26, ci = 0.11, ri = 0.54]
154. Male — Oblique metapleural suture (apterous/brachypterous/obliterated = -): 0 = Running anteroventrally from endophragmal pit; 1 = Running horizontally from endophragmal pit; 2 = Running anterodorsally from endophragmal pit. [11%; length = 22, ci = 0.09, ri = 0.41]
155. Male — Mesosternum, just anterior to mesocoxae: 0 = Smoothly rounded; 1 = With paired transverse/oblique carinae (may be toothed mesally); 2 = With paired lamellate projections mesally. (NONADDITIVE) [6%; length = 18, ci = 0.11, ri = 0.33]
156. Male — Mesosternum, midway to anterior margin: 0 = Evenly convex; 1 = With distinct paired teeth or tubercles; 2 = With distinct paired (separated) transverse carinae or ridges; 3 = With paired longitudinal high lamellae acuminate apically. (NONADDITIVE) [6%; length = 15, ci = 0.20, ri = 0.36]
157. Male — Mesocoxae, contiguity: 0 = Contiguous mesally; 1 = Slightly separated mesally. [0%; length = 3, ci = 0.33, ri = 0.66]
158. Male — Mesocoxa, insertion: 0 = Large basicoxite, coxal cavities large and approximated; 1 = Large basicoxite, coxal cavities large and widely separated; 2 = Reduced basicoxite, coxal cavities small and widely separated. [0%; length = 3, ci = 0.66, ri = 0.50]
159. Male — Metasternum, posterior median process (absent = -): 0 = Shorter than coxal height, tridentate; 1 = Shorter than coxal height, shallowly bidentate; 2 = Shorter than coxal height, deeply bidentate; 3 = Shorter than coxal height, unidentate; 4 = Longer than coxal height, acutely unidentate; 5 = Longer than coxal height, obtusely unidentate; 6 = Shorter than coxal height, tridentate with median tooth incised; 7 = Longer than coxal height, tridentate with median tooth acute; 8 = Shorter than coxal height, forming a transverse crenulate ridge. (NONADDITIVE) [16%; length = 29, ci = 0.27, ri = 0.54]
160. Male — Metacoxal cavities: 0 = Open; 1 = Partially closed; 2 = Closed. [0%; length = 3, ci = 0.66, ri = 0.90]
161. Male — Tarsal claws: 0 = Midventrally toothed; 1 = Simple; 2 = Ventrally basally lamellate, distinct apex acute; 3 = Apically deeply bifid. (NONADDITIVE) [0%; length = 3, ci = 1.00, ri = 1.00]
162. Male — Fore tibia, inner (anterior) secretory structure: 0 = None; 1 = Broad coarsely setose delimited patch; 2 = Linear to oval finely perforated depression; 3 = Vertically elongate groove/pore; 4 = Obliquely elongate groove/pore; 5 = Obliquely oval to circular pore; 6 = Basal elongate/oval and separated apical round pores. (NONADDITIVE) [12%; length = 26, ci = 0.23, ri = 0.54]
163. Male — Fore tibia, outer (posterior) secretory structure: 0 = None; 1 = Linear to oval finely perforated depression; 2 = Obliquely elongate groove/pore; 3 = Obliquely oval to circular pore. (NONADDITIVE) [7%; length = 12, ci = 0.16, ri = 0.37]
164. Male — Fore calcar blade: 0 = Linearly narrow; 1 = Expanded, longish >0.5 × length of calcar; 2 = Expanded, almost square, <0.4 length of calcar. (NONADDITIVE) [0%; length = 3, ci = 0.66, ri = 0.93]
165. Male — Fore tarsomeres, apicoventral median ovoid pulvillus: 0 = Absent; 1 = On 4th tarsomere only; 2 = On 3rd & 4th tarsomeres; 3 = On 2nd–4th tarsomeres; 4 = On 1st–4th tarsomeres. [5%; length = 14, ci = 0.28, ri = 0.68]
166. Male — Meso- and metatibial articulated spines, mean number: 0 = 0–4; 1 = 5–9; 2 = 10–14; 3 = 15–19; 4 = 20–24; 5 = >24. [7%; length = 25, ci = 0.20, ri = 0.52]
167. Male — Mesotarsomeres, apicoventral median ovoid pulvillus: 0 = Absent; 1 = On 4th tarsomere only; 2 = On 3rd & 4th tarsomeres; 3 = On 2nd–4th tarsomeres; 4 = On 1st–4th tarsomeres. [6%; length = 14, ci = 0.28, ri = 0.54]
168. Male — Metacoxa, mesally: 0 = Simple; 1 = Longitudinally carinate; 2 = Dentate; 3 = With setaceous pit. (NONADDITIVE) [8%; length = 9, ci = 0.22, ri = 0.79]
169. Male — Metatibia, posterior longitudinal smooth glabrous ridge/carina: 0 = Absent; 1 = Present. [0%; length = 2, ci = 0.50, ri = 0.93]
170. Male — Metatibia, apex dorsally: 0 = Evenly rounded; 1 = With elevated tubercle bearing spine(s); 2 = With distinct cylindrical process bearing spine. [4%; length = 2, ci = 0.50, ri = 0.00]
171. Male — Metatibia, posteroapical secretory structure: 0 = Absent; 1 = Present, delimited patch of dense setae; 2 = Present, a small pore; 3 = Present, a deep narrow longitudinal groove. (NONADDITIVE) [2%; length = 8, ci = 0.37, ri = 0.73]
172. Male — Metatibia, apical spurs: 0 = Both unmodified; 1 = Inner modified as cleaner. [0%; length = 1, uninformative]
173. Male — Metatarsomeres, apicoventral median ovoid pulvillus: 0 = Absent; 1 = On 4th tarsomere only; 2 = On 3rd & 4th tarsomeres; 3 = On 2nd–4th tarsomeres; 4 = On 1st–4th tarsomeres. [7%; length = 12, ci = 0.41, ri = 0.50]
174. Male — Wings and tegula: 0 = Fully developed; 1 = Brachypterous, wing exceeding propodeum apex, tegula present; 2 = Micropterous, wing shorter than propodeum base, tegula present; 3 = Apterous, tegula absent. [2%; length = 16, ci = 0.18, ri = 0.18]
175. Male — Fore wing, extent of venation (apterous/brachypterous = -): 0 = Reaching distal margin; 1 = Ending before distal margin. [0%; length = 1, ci = 1.00, ri = 1.00]
176. Male — Fore wing, vein Sc+R (apterous/brachypterous = -): 0 = <0.5 × length of 1st abscissa of RS; 1 = Subequal to 1st abscissa of RS; 2 = >1.5 × length of 1st abscissa of RS. [6%; length = 16, ci = 0.12, ri = 0.30]
177. Male — Fore wing, pterostigma, sclerotization (apterous/brachypterous = -): 0 = Entirely well sclerotized; 1 = Sclerotization reduced anteriorly; 2 = Unsclerotized; 3 = Entirely faintly sclerotized. (NONADDITIVE) [2%; length = 8, ci = 0.37, ri = 0.87]
178. Male — Fore wing, pterostigma, delimitation (apterous/brachypterous = -): 0 = Completely delimited by distinct veins or completely sclerotized; 1 = Vein SC lost or much reduced, pterostigma not delimited basally. [0%; length = 5, ci = 0.20, ri = 0.60]
179. Male — Fore wing, pterostigma, base (apterous/brachypterous = -): 0 = With interruption/constriction in C and Sc+R; 1 = With interruption/constriction in Sc+R only; 2 = Without interruptions/constrictions. [2%; length = 17, ci = 0.11, ri = 0.71]
180. Male — Fore wing, pterostigma, shape (apterous/brachypterous = -): 0 = Elongate, broader than base; 1 = Elongate, as narrow as base; 2 = Short, broader than base; 3 = Short, as narrow as base; 4 = Very short, narrowed from base; 5 = Minuscule or absent. (NONADDITIVE) [5%; length = 22, ci = 0.22, ri = 0.66]
181. Male — Fore wing, radial (marginal) cell apex (apterous/brachypterous = -): 0 = Acute; 1 = Blunt; 2 = Rounded; 3 = Obtuse with posterior spur. (NONADDITIVE) [10%; length = 24, ci = 0.12, ri = 0.27]
182. Male — Fore wing, vein RS2 (apterous/brachypterous = -): 0 = Absent; 1 = Present and complete, basally tubular or solid nebulous; 2 = Present and complete, entirely pigmented spectral; 3 = Apically present but basally absent, pigmented nebulous or spectral; 4 = Present as short stub only. (NONADDITIVE) [14%; length = 12, ci = 0.33, ri = 0.66]
183. Male — Fore wing, closed submarginal cells (apterous/brachypterous = -): 0 = Three, all veins tubular; 1 = Three, vein 3r-m nebulous; 2 = Two, all veins tubular; 3 = Two, vein 2r-m nebulous; 4 = One. [15%; length = 13, ci = 0.30, ri = 0.74]
184. Male — Fore wing, cell 1R1 (first submarginal) (apterous/brachypterous = -): 0 = Rudiment of crossvein 1r-rs present, at least with third abscissa of vein RS slightly thickened near base; 1 = Rudiment of crossvein 1r-rs absent, third abscissa of vein RS of even width throughout. [2%; length = 3, ci = 0.33, ri = 0.50]
185. Male — Fore wing, vein RS third abscissa, bulla (apterous/brachypterous = -): 0 = Present, even if indistinct; 1 = Absent. [2%; length = 6, ci = 0.16, ri = 0.88]
186. Male — Fore wing, vein RS third abscissa, course (apterous/brachypterous = -): 0 = With distinct angle; 1 = With weak angle; 2 = Straight or very weakly and evenly curved. [10%; length = 24, ci = 0.08, ri = 0.56]
187. Male — Fore wing, cell 1S (second submarginal) (apterous/brachypterous = -): 0 = Sessile anteriorly; 1 = Petiolate anteriorly. [1%; length = 1, ci = 1.00, ri = 1.00]
188. Male — Fore wing, crossvein 3r-m (absent/apterous/brachypterous = -): 0 = With bulla; 1 = Without bulla. [4%; length = 5, ci = 0.20, ri = 0.87]
189. Male — Fore wing, jugal lobe (apterous/brachypterous = -): 0 = Present; 1 = Absent. [0%; length = 1, ci = 1.00, ri = 1.00]
190. Male — Hind wing, basal hamuli, occurrence (apterous/brachypterous = -): 0 = Present; 1 = Absent. [0%; length = 1, ci = 1.00, ri = 1.00]
191. Male — Hind wing, basal hamuli, position (none/apterous/brachypterous = -): 0 = Dispersed; 1 = Basal cluster. [0%; length = 3, ci = 0.33, ri = 0.60]
192. Male — Hind wing, apical hamuli (apterous/brachypterous = -): 0 = <11; 1 = >10. [4%; length = 8, ci = 0.12, ri = 0.41]
193. Male — Hind wing, vein RS junction with vein SC (apterous/brachypterous = -): 0 = At acute angle; 1 = At right angle. [9%; length = 16, ci = 0.06, ri = 0.53]
194. Male — Hind wing, crossvein r-m (apterous/brachypterous = -): 0 = Distal; 1 = Proximal, complete; 2 = Proximal, incomplete; 3 = Absent. [11%; length = 16, ci = 0.18, ri = 0.69]
195. Male — Hind wing, vein M free apical section (apterous/brachypterous = -): 0 = Present; 1 = Absent. [8%; length = 11, ci = 0.09, ri = 0.47]
196. Male — Hind wing, vein Cu free apex (apterous/brachypterous = -): 0 = Present, even if only a small stub or nebulous trace; 1 = Absent. [1%; length = 3, ci = 0.33, ri = 0.75]
197. Male — Hind wing, crossvein cu-a (apterous/brachypterous = -): 0 = Present, tubular or solid; 1 = Present, nebulous; 2 = Absent. [9%; length = 24, ci = 0.08, ri = 0.63]
198. Male — Hind wing, vein A free apical section (apterous/brachypterous = -): 0 = Present; 1 = Absent. [1%; length = 4, ci = 0.25, ri = 0.57]
199. Male — Hind wing, anal lobe (apterous/brachypterous = -): 0 = Moderate incision on margin; 1 = Shallow definite notch on margin; 2 = Not indicated on margin (at most very shallowly sinuate). [0%; length = 2, ci = 1.00, ri = 1.00]
200. Male — Hind wing, jugal lobe (apterous/brachypterous = -): 0 = Present, large with incision about half length; 1 = Present, small with incision nearly to base; 2 = Absent. [0%; length = 2, ci = 1.00, ri = 1.00]
201. Male — Tergum I and propodeum pubescence: 0 = Entirely simple; 1 = Some brachyplumose; 2 = Some fully plumose. [3%; length = 6, ci = 0.33, ri = 0.87]
202. Male — Tergum I, shape: 0 = Gradually broadened posteriorly, ≥0.5 × length tergum II, apically sessile on tergum II; 1 = Gradually broadened posteriorly, ≥0.5 × length tergum II, apically constricted from tergum II; 2 = Gradually broadened posteriorly, <0.5 × length tergum II, apically sessile on tergum II; 3 = Gradually broadened posteriorly, <0.5 × length tergum II, apically constricted from tergum II; 4 = Parallel-sided posteriorly, ≥0.4 × length tergum II, discontinuous with tergum II; 5 = Entirely parallel-sided, <0.5 × length tergum II, discontinuous with tergum II. (NONADDITIVE) [9%; length = 17, ci = 0.29, ri = 0.68]
203. Male — Tergum I, apical width: 0 = >0.75 × width tergum II; 1 = >0.5 <0.75 × width tergum II; 2 = <0.5 × width tergum II. [3%; length = 18, ci = 0.11, ri = 0.40]
204. Male — Tergum 1, anterodorsal profile: 0 = Broadly convex; 1 = Anterior and dorsal faces merging; 2 = Anterior and dorsal faces distinct. [9%; length = 22, ci = 0.09, ri = 0.53]
205. Male — Tergum II, felt line, presence: 0 = Absent; 1 = Present. [2%; length = 5, ci = 0.20, ri = 0.71]
206. Male — Tergum II, felt line, form (absent = -): 0 = Dispersed traces; 1 = Linear, superficial; 2 = Linear, abruptly invaginated. (NONADDITIVE) [0%; length = 4, ci = 0.50, ri = 0.66]
207. Male — Tergum II, apical fringe: 0 = Setae many, slender arcuate, simple to slightly flattened; 1 = Setae many, some densely plumose; 2 = Setae many, apically split; 3 = Setae many, strong and curved; 4 = Setae few, strong, long, convergent. (NONADDITIVE) [1%; length = 6, ci = 0.50, ri = 0.70]
208. Male — Tergum III, stridulitrum: 0 = Absent; 1 = Present. [0%; length = 3, ci = 0.33, ri = 0.60]
209. Male — Sternum I, differentiation: 0 = Smoothly overlapping sternum II; 1 = Briefly declivous and abutting sternum II; 2 = Depressed posteriorly, constricted and abutting sternum II. [1%; length = 2, ci = 1.00, ri = 1.00]
210. Male — Sternum II, lateral felt line, presence: 0 = Absent; 1 = Present. [7%; length = 18, ci = 0.05, ri = 0.50]
211. Male — Sternum II, lateral felt line, form (absent = -): 0 = Dispersed traces only; 1 = Distinct but minute; 2 = Well developed. [5%; length = 15, ci = 0.13, ri = 0.27]
212. Male — Sternum VII: 0 = Entirely exposed, about as long as sternum VI; 1 = Partly exposed, much shorter than sternum VI; 2 = Concealed. [4%; length = 22, ci = 0.09, ri = 0.58]
213. Male — Hypopygium, visibility: 0 = Almost entirely exposed, lateral margin entire or only shallowly notched; 1 = Almost entirely concealed, lateral margin very deeply incised, hypopygium tri- or pentalobate. [0%; length = 1, ci = 1.00, ri = 1.00]
214. Male — Hypopygium, exposed surface (hidden = -): 0 = Convex to flat, more or less evenly sculptured, punctate to smooth; 1 = Concave, more or less evenly punctured to smooth with lateral longitudinal carina; 2 = Evenly convex, with median tubercle on basal half; 3 = Convex mediolongitudinally, with abrupt lateral depression; 4 = Convex, with median smooth ridge; 5 = Convex to flat, with sublateral paired longitudinal oblique ridges; 6 = Convex, with median Y-shaped ridge; 7 = With longitudinal smooth median depression, dense punctures laterally; 8 = With median excavation, lateral peg-like projection. (NONADDITIVE) [5%; length = 18, ci = 0.38, ri = 0.47]
215. Male — Hypopygium, apex: 0 = Simple, rounded or obtuse; 1 = With shallow broad median emargination; 2 = With simple deep narrow median emargination; 3 = With broad lobed median emargination; 4 = With deep narrow median emargination with internal sclerites; 5 = With median tooth or peg; 6 = With two small approximated teeth or slight notch; 7 = Broadly bilobed; 8 = With two small lateral teeth; 9 = With two separated moderate teeth. (NONADDITIVE) [4%; length = 20, ci = 0.45, ri = 0.38]
216. Male — Cercus: 0 = Present; 1 = Absent. [0%; length = 3, ci = 0.33, ri = 0.00]
217. Male — Cercus, form (absent = -): 0 = Elongate, cylindrical or weakly evenly broadened apically; 1 = Elongate, strongly clavate (with narrow basal stalk); 2 = Elongate, narrow, flattened; 3 = Short, base narrow, distinctly flattened; 4 = Short, base widened, apex narrowed, distinctly flattened; 5 = Short, flattened basally, clavate apically; 6 = Short, evenly clavate; 7 = Vestigial; 8 = Broad-based diskiform, flattened. (NONADDITIVE) [0%; length = 24, ci = 0.33, ri = 0.62]
218. Male — Gonobase, form: 0 = Complete, dorsal and ventral lengths similar, as long as paramere base; 1 = Complete, dorsal and ventral lengths similar, very short and annular; 2 = Complete, dorsal length shorter than ventral, ventrally as long as paramere base; 3 = Complete, dorsal and ventral lengths similar, longer than paramere base; 4 = Complete, dorsal length shorter than ventral, ventrally longer than paramere base; 5 = Complete, dorsal length shorter than ventral, much shorter than paramere base; 6 = Dorsally incomplete, dorsal length shorter than ventral, ventrally as long as paramere base; 7 = Dorsally absent, very short. (NONADDITIVE) [2%; length = 25, ci = 0.28, ri = 0.68]
219. Male — Gonostylus, form, lateral view: 0 = Short, lamellate with rounded apex; 1 = Short, tapered with narrow to acute apex; 2 = Elongate, tapered with acute apex; 3 = Elongate, lamellate with rounded apex. (NONADDITIVE) [2%; length = 9, ci = 0.33, ri = 0.68]
220. Male — Gonostylus, apical curvature, lateral view: 0 = Upcurved; 1 = Straight; 2 = Downcurved. [4%; length = 11, ci = 0.18, ri = 0.87]
221. Male — Gonostylus, dorsal transverse suture (distant from gonobase): 0 = Well developed, extending at least halfway to lateral margin; 1 = Absent or short, longitudinal suture ending distant from gonobase; 2 = Absent, longitudinal suture reaching gonobase. [3%; length = 25, ci = 0.08, ri = 0.63]
222. Male — Gonostylus, parapenial lobe: 0 = Absent; 1 = Present. [1%; length = 2, ci = 0.50, ri = 0.83]
223. Male — Gonostylus, dorsal oblique stout setae: 0 = None; 1 = Present, arising under dorsal flange. [0%; length = 1, ci = 1.00, ri = 1.00]
224. Male — Gonapophysis IX (penis valve), fusion: 0 = Fused dorsally for most of length; 1 = Free for most of length. [0%; length = 1, ci = 1.00, ri = 1.00]
225. Male — Gonapophysis IX (penis valve), shape: 0 = Apex elongate, rounded, no ventral tooth; 1 = Apex rounded, ventral tooth about midway; 2 = Apex dorsally produced, ventral tooth about midway; 3 = Apex dorsally simple, ventral tooth on apical half; 4 = Apex rounded, produced, ventral prominence about midway. (NONADDITIVE) [0%; length = 4, ci = 1.00, ri = 1.00]
226. Male – Gonapophysis IX (penis valve), articulated spines or long setae: 0 = Absent; 1 = Present, strong short spines; 2 = Present, thick long setae. (NONADDITIVE) [0%; length = 2, uninformative]
227. Male — Gonapophysis IX (penis valve), right: 0 = Same shape and length as left gonapophysis IX; 1 = Longer and more elaborate than left gonapophysis IX. [1%; length = 3, ci = 0.33, ri = 0.71]
228. Male — Volsella, basal lobe: 0 = Present, as distinct prominent inner projection; 1 = Present, as rounded ventral long-setose expansion well differentiated from slender apicodorsal section; 2 = Absent, even though slight inner swelling may be evident or base may be somewhat broader than apex. (NONADDITIVE) [0%; length = 10, ci = 0.20, ri = 0.27]
229. Male — Volsella, digitus: 0 = Present, distinct; 1 = Absent or scarcely discernible. [0%; length = 3, ci = 0.33, ri = 0.75]
230. Male — Volsella, paracuspis: 0 = Absent; 1 = Present, as tubercle/swelling/projection at base of cuspis and lateral to digitus. [0%; length = 10, ci = 0.10, ri = 0.75]
Data matrix for phylogenetic analysis of sub/genera of Mutillidae and four outgroup taxa
Polymorphisms are indicated between square brackets, inapplicable characters are indicated by hyphens, and missing data are indicated by question marks. (An operational version in Nona format is supplied as Suppl. material
Proposed higher classification of genera and subgenera of Mutillidae
All currently valid genera (216) and subgenera (30) are listed (for convenience simply under the heading of “Genera”), indicating the sexes known for each (whether described or not), and those included in the current analysis are in boldface. Details for each name appear in
Family: Mutillidae Latreille, 1802
Subfamily: Myrmosinae Fox, 1894
Tribe: Kudakrumiini Krombein, 1979
Genera: Kudakrumia Krombein, 1979 (♂, ♀); Leiomyrmosa Wasbauer, 1973 (♀); Myrmosula Bradley, 1917 (♂, ♀); Nothomyrmosa Krombein, 1979 (♀); Protomutilla† Bischoff, 1916 (♂, ♀); Pseudomyrmosa Suárez, 1980 (♂, ♀)
Tribe: Myrmosini Fox, 1894
Genera: Carinomyrmosa Lelej, 1981 (♂, ♀); Erimyrmosa Lelej, 1984b (♂); Krombeinella Pate, 1947 (♂, ♀); Myrmosa Latreille, 1797 (♂, ♀); Myrmosina Krombein, 1940 (♂); Paramyrmosa Saussure, 1880 (♂, ♀); Taimyrmosa Lelej, 2005 (♂, ♀)
Subfamily: Pseudophotopsidinae Bischoff, 1920
Genus: Pseudophotopsis André, 1896 (♂, ♀)
Subfamily: Ticoplinae Nagy, 1970
Tribe: Smicromyrmillini Argaman, 1988
Genera: Cameronilla Lelej in Lelej & Krombein, 2001 (♂); Eosmicromyrmilla Lelej & Krombein, 2001 (♂, ♀); Hindustanilla Lelej in Lelej & Krombein, 2001 (♂, ♀); Smicromyrmilla Suárez, 1965 (♂, ♀)
Tribe: Ticoplini Nagy, 1970
Genera: Areotilla Bischoff, 1920 (♂, ♀); Nanomutilla André, 1900 (♂, ♀)
Subfamily: Rhopalomutillinae Schuster, 1949
Genera: Bischoffiella Brothers & Nonveiller in Brothers, 2015 (♂, ♀); Pherotilla Brothers, 2015 (♂, ♀); Rhopalomutilla André, 1901 (♂, ♀); Rimulotilla Brothers, 2015 (♂, ♀)
Subfamily: Sphaeropthalminae Schuster, 1949 (1903)
Tribe: Sphaeropthalmini Schuster, 1949 (1903)
Genera: Acanthophotopsis Schuster, 1958 (♂); Acrophotopsis Schuster, 1958 (♂); Allotilla Schuster, 1949 (♂, ♀); Ceratophotopsis Schuster, 1949 (♂); Chilemutilla Cambra & Quintero, 2007 (♂, ♀); Chilephotopsis Cambra & Quintero, 2006 (♂); Cystomutilla André, 1896 (♂, ♀); Dilophotopsis Schuster, 1958 (♂, ♀); Hemutilla Lelej, Tu & Chen in Tu et al., 2014 (♂, ♀); Laminatilla Pitts, 2007 (♂); Limaytilla Casal, 1964 (♂, ♀); Morsyma Fox, 1899 (♂, ♀); Nanotopsis Schuster, 1949 (♂, ♀); Odontophotopsis Viereck, 1903 (♂, ♀); Photomorphina Schuster, 1952 (♂, ♀); Photomorphus Viereck, 1903 (♂, ♀); Ptilomutilla André, 1905 (♀); Scaptodactyla Burmeister, 1875 (♂, ♀); Schusterphotopsis Pitts, 2003 (♂); Sphaeropthalma Blake, 1871 (♂, ♀); Stethophotopsis Pitts in Pitts & McHugh, 2000 (♂, ♀); Tallium André, 1902 (♂, ♀); Xenomorphus Schuster, 1958 (♂); Xystromutilla André, 1905 (♂, ♀)
Tribe: Dasymutillini Brothers & Lelej, trib. n.
Genera: Ancistrotilla Brothers, 2012 (♂, ♀); Ascetotilla Brothers, 1971 (♂, ♀); Australotilla Lelej, 1983 (♂, ♀); Bothriomutilla Ashmead, 1899 (♂, ♀); Cephalomutilla André, 1908 (♂, ♀); Dasymutilla Ashmead, 1899 (♂, ♀); Ephutomorpha André, 1902 (♂, ♀); Eurymutilla Ashmead, 1899 (♀); [Eurymutilla (genus near this) (♂, ♀);] Frigitilla Williams in Bartholomay et al., 2015 (♂, ♀); Gogoltilla Williams, Brothers & Pitts, 2011 (♂, ♀); Leucospilomutilla Ashmead, 1903 (♂, ♀); Lomachaeta Mickel, 1936 (♂, ♀); Neomutilla Reed, 1898 (♂, ♀); Odontomyrme Lelej, 1983 (♂, ♀); Ponerotilla Brothers, 1994 (♀); Protophotopsiella Schuster, 1949 (♂, ♀); Protophotopsis Schuster, 1947 (♂, ♀); Reedomutilla Mickel, 1964 (♂, ♀); Suareztilla Casal, 1968 (♂, ♀); Tobantilla Casal, 1965 (♂, ♀); Traumatomutilla André, 1901 (♂, ♀);
Tribe: Pseudomethocini Brothers, 1975
Subtribe: Euspinoliina Brothers & Lelej, subtrib. n.
Genera: Atillum André, 1902 (♂, ♀); Euspinolia Ashmead, 1903 (♂, ♀); Hoplocrates Mickel, 1937 (♂, ♀)
Subtribe: Pseudomethocina Brothers, 1975
Genera: Anomophotopsis Schuster, 1949 (♂, ♀); Calomutilla Mickel, 1952 (♂, ♀); Chaetotilla Schuster, 1949 (♂); Darditilla Casal, 1965 (♂, ♀); Dimorphomutilla Ashmead, 1903 (♂, ♀); Gurisita Casal, 1970 (♀); Hoplognathoca Suárez, 1962 (♂, ♀); Hoplomutilla Ashmead, 1899 (♂, ♀); Horcomutilla Casal, 1962 (♂, ♀); Invreiella Suárez, 1966 (♀); Lophomutilla Mickel, 1952 (♂, ♀); Lophostigma Mickel, 1952 (♂, ♀); Lynchiatilla Casal, 1963 (♂, ♀); Mickelia Suárez, 1966 (♀); Myrmilloides André, 1902 (♂, ♀); Pappognatha Mickel, 1939 (♂, ♀); Patquiatilla Casal, 1962 (♂, ♀); Pertyella Mickel, 1952 (♂, ♀); Pseudomethoca Ashmead, 1896 (♂, ♀); Seabratilla Casal, 1963 (♀); Vianatilla Casal, 1962 (♂, ♀)
Subfamily: Dasylabrinae Invrea, 1964
Tribe: Apteromutillini Brothers & Lelej, trib. n.
Genera: Apteromutilla Ashmead, 1903 (♂, ♀); Brachymutilla André, 1901 (♂, ♀); Liotilla Bischoff, 1920 (♂, ♀)
Tribe: Dasylabrini Invrea, 1964
Genera: Baltilla Lelej, 1976 (♂, ♀); Chrestomutilla Brothers, 1971 (♂, ♀); Craspedopyga Lelej, 1976 (♂, ♀); Dasylabris Radoszkowski, 1885 (♂, ♀); Dasylabroides André, 1901 (♂, ♀); Inbaltilla Lelej, 1976 (♂, ♀); Jaxartilla Lelej, 1984 (♂); Orientilla Lelej, 1979 (♂, ♀); Seyrigilla Krombein, 1972 (♂, ♀); Stenomutilla André, 1896 (♂, ♀); Tricholabiodes Radoszkowski, 1885 (♂, ♀)
Subfamily: Myrmillinae Bischoff, 1920
Genera: Arnoldtilla Nonveiller, 1996 (♂, ♀); Bethsmyrmilla Krombein & Lelej, 1999 (♀); Bidecoloratilla Turrisi & Matteini Palmerini in Turrisi et al., 2015 (♂, ♀); Bimaculatilla Turrisi & Matteini Palmerini in Turrisi et al., 2015 (♂, ♀); Bischoffitilla Lelej, 2002 (♂, ♀); Bisulcotilla Bischoff, 1920 (♂); Blakeius Ashmead, 1903 (♂, ♀); Botswanotilla Nonveiller, 1996 (♂); Brahmatilla Lelej, 2005 (♀); Cataractaetilla Nonveiller, 1996 (♂, ♀); Ceratotilla Bischoff, 1920 (♂, ♀); Clinotilla Arnold, 1956 (?♂, ♀); Eurygnathilla Skorikov, 1927 (♂, ♀); Labidomilla André, 1902 (♂, ♀); Liomutilla André, 1907 (♂, ♀); Myrmilla Wesmael, 1851 (♂, ♀); Myrmotilla Bischoff, 1920 (♀); Odontotilla Bischoff, 1920 (♂, ♀); Odontotilloides Nonveiller, 1996 (♂, ♀); Omotilla Invrea, 1943 (♂, ♀); Platymyrmilla André, 1900 (♂, ♀); Pseudomutilla Costa, 1885 (♂, ♀); Pygomilla Hammer, 1955 (♀); Saganotilla Invrea, 1943 (♂, ♀); Sigilla Skorikov, 1927 (♂, ♀); Somaliatilla Nonveiller, 1996 (♀); Spilomutilla Ashmead, 1903 (♂, ♀); Squamulotilla Bischoff, 1920 (♂); Viereckia Ashmead, 1903 (♂, ♀)
Subfamily: Mutillinae Latreille, 1802
Tribe: Ctenotillini Brothers & Lelej, trib. n.
Genera: Arcuatotilla Nonveiller, 1998 (♂, ♀); Bidentotilla Nonveiller, 1979 (♂); Cephalotilla Bischoff, 1920 (♂, ♀); Chaetomutilla Nonveiller, 1979 (♂, ♀); Ctenotilla Bischoff, 1920 (♂, ♀); Lehritilla Lelej, 2005 (♂); Mimecomutilla Ashmead, 1903 (♂, ♀); Mimecotilla Nonveiller, 1998 (♂, ♀); Montanomutilla Nonveiller, 1979 (♀); Pristomutilla Ashmead, 1903 (♂, ♀); Strangulotilla Nonveiller, 1979 (♂, ♀); Taeniotilla Nonveiller, 1979 (♂); Zeugomutilla Chen, 1957 (♂, ♀)
Tribe: Smicromyrmini Bischoff, 1920
Genera: Andreimyrme Lelej, 1995 (♂, ♀); Antennotilla Bischoff, 1920 (♂); Astomyrme Schwartz, 1984 (♂, ♀); Corytilla Arnold, 1956 (♂, ♀); Ctenoceraea Nonveiller, 1993 (♂); Dentilla Lelej in Lelej & Kabakov, 1980 (♂, ♀); Ephucilla Lelej 1995 (♂, ♀); Ephutomma Ashmead, 1899 (♂, ♀); Eremotilla Lelej, 1985 (♂, ♀); Erimyrme Lelej, 1985 (♂, ♀); Guineomutilla Suárez, 1977 (♀); Gynandrotilla Arnold, 1946 (♂); Indratilla Lelej, 1993 (♂, ♀); Karunaratnea Lelej, 2005 (♂, ♀); Mickelomyrme Lelej, 1995 (♂, ♀); Nemka Lelej, 1985 (♂, ♀); Nordeniella Lelej, 2005 (♂, ♀); Nuristanilla Lelej in Lelej & Kabakov, 1980 (♀); Paglianotilla Lelej in Lelej & van Harten, 2006 (♂); Physetopoda Schuster, 1949 (♂, ♀); Promecilla André, 1902 (♂, ♀); Psammotherma Latreille, 1825 (♂); Pseudocephalotilla Bischoff, 1920 (♂, ♀); Rasnitsynitilla Lelej in Lelej & van Harten, 2006 (♂); Rhombotilla Nagy, 1966 (♀); Sinotilla Lelej, 1995 (♂, ♀); Skorikovia Ovtchinnikov, 2002 (♂, ♀); Smicromyrme Thomson, 1870 (♂, ♀); Sulcotilla Bischoff, 1920 (♂, ♀); Tsunekimyrme Lelej, 1995 (♂)
Tribe: Mutillini Latreille, 1802
Subtribe: Ephutina Ashmead, 1903 (= Odontomutillini Lelej, 1983, syn. n.)
Genera: Cockerellidia Lelej & Krombein, 1999 (♀); Ephuamelia Casal, 1968 (♂); Ephuchaya Casal, 1968 (♂); Ephuseabra Casal, 1968 (♂); Ephusuarezia Casal, 1968 (♂); Ephuta Say, 1836 (♂, ♀); Ephutopsis Ashmead, 1904 (♂, ♀); Karlidia Lelej in Lelej & Krombein, 1999 (♀); Odontomutilla Ashmead, 1899 (♂, ♀); Onoretilla Pagliano in Pagliano, Cambra & Quintero, 2017 (♂); Xenochile Schuster, 1957 (♂); Yamanetilla Lelej, 1996 (♂, ♀)
Subtribe: Mutillina Latreille, 1802
Genera: Barymutilla André, 1901 (♂, ♀); Hadrotilla Bischoff, 1920 (♂, ♀); Kurzenkotilla Lelej, 2005 (♀); Macromyrme Lelej, 1984 (♀); Mutilla Linnaeus, 1758 (♂, ♀); Nanomyrme Lelej, 1977 (♀); Ronisia Costa, 1858 (♂, ♀); Standfussidia Lelej, 2005 (♀); Storozhenkotilla Lelej, 2005 (♂, ♀); Tropidotilla Bischoff, 1920 (♂, ♀)
Tribe: Trogaspidiini Bischoff, 1920 (= Petersenidiina Lelej, 1996, syn. n.)
Genera: Acanthomutilla Nonveiller, 1995 (♂, ♀); Acutitropidia Nonveiller, 1995 (♂, ♀); Allotropidia Nonveiller, 1996 (♂); Amblotropidia Nonveiller, 1995 (♂, ♀); Arcuatotropidia Nonveiller, 1995 (♂); Artiotilla Invrea, 1950 (♂, ♀); Aureotilla Bischoff, 1920 (♂, ♀); Carinotilla Nonveiller, 1973 (♂, ♀); Chilotropidia Nonveiller, 1995 (♂, ♀); Chrysotilla Bischoff, 1920 (♂, ♀); Curvitropidia Nonveiller, 1995 (♂, ♀); Dentotilla Nonveiller, 1977 (♂, ♀); Diacanthotilla Nonveiller, 1995 (♀); Dolichomutilla Ashmead, 1899 (♂, ♀); Eotrogaspidia Lelej, 1996 (♂, ♀); Glossotilla Bischoff, 1920 (♂, ♀); Hildbrandetia Özdikmen, 2005 (♀); Inflatispidia Nonveiller, 1995 (♂, ♀); Karlissaidia Lelej, 2005 (♂, ♀); Krombeinidia Lelej, 1996 (♂, ♀); Lobotilla Bischoff, 1920 (♂, ♀); Lobotropidia Nonveiller, 1995 (♂, ♀); Lophotilla Bischoff, 1920 (♂); Neotrogaspidia Lelej, 1996 (♂, ♀); Nonveilleridia Lelej, 1996 (♂); Orientidia Lelej, 1996 (♂, ♀); Pagdenidia Lelej, 1996 (♂, ♀); Petersenidia Lelej in Lelej & Yamane, 1992 (♂, ♀); Promecidia Lelej, 1996 (♂, ♀); Protrogaspidia Lelej, 1996 (♂); Pseudolophotilla Nonveiller & Ćetković, 1995 (♂, ♀); Radoszkowskitilla Lelej, 2005 (♂, ♀); Serendibiella Lelej, 2005 (♂); Seriatospidia Nonveiller & Ćetković, 1996 (♀); Spinulomutilla Nonveiller, 1994 (♂, ♀); Spinulotilla Bischoff, 1920 (♂, ♀); Sylvotilla Viette, 1978 (♀); Taiwanomyrme Tsuneki, 1993 (♂, ♀); Timulla Ashmead, 1899 (♂, ♀); Trispilotilla Bischoff, 1920 (♂, ♀); TrogaspidiaAshmead, 1899 (♂, ♀); Tuberocoxotilla Nonveiller, 1980 (♂); Vanhartenidia Lelej in Lelej & van Harten, 2006 (♂, ♀); Wallacidia Lelej & Brothers, 2008 (♂, ♀); Zavatilla Tsuneki, 1993 (♂)
Family Mutillidaeincertae sedis
Genus: Cretavus† Sharov, 1957 (♂)
Data matrix for phylogenetic analysis of sub/genera of Mutillidae and four outgroup taxa
Data type: Taxon versus character-state matrix
Explanation note: This is an operational version of the data matrix in Nona format.