Corresponding author: Denis J. Brothers (
Academic editor: Michael Engel
The taxonomic placement of an enigmatic species of wasp known from two specimens in Late Cretaceous New Jersey amber is investigated through cladistic analyses of 90 morphological characters for 33 terminals ranging across non-Aculeata, non-
The phylogeny of the
As chrysidoids, plumariids are very unusual morphologically, the males having broad wings with a relatively rich wing venation including well developed accessory veins in the apical membrane similar to those of many of the very distantly related
Previous estimates of relationships.
Within the
No fossil
The two amber pieces (
Previous cladistic analyses of the
Parsimony analyses by TNT (
Male. Pronotum forming a short convex band reaching tegula; propleura closely associated, anterodorsally exposed as a short neck, posteriorly swollen and transversely truncate; prosternum short and scarcely exposed medially; mesopleuron large and swollen; metasternum somewhat depressed. Forewing with pterostigma very large, seven closed cells (costal, basal, subbasal, marginal, first and second submarginals, first discal), second submarginal cell with long anterior margin, no accessory veins in apical membrane. Hind wing with closed cells (basal and subbasal at least), anal (vannal/plical) lobe well developed; jugal lobe absent. Coxae subglobose, trochanters inserted apically.
Female. Unknown.
The genus name, which is masculine, is derived from “
Male. Compound eye oval with convex inner margin; antenna with many long fine erect setae (number of antennomeres unknown); mandible with four apical teeth along truncate apical margin; maxillary palp at least 5-segmented; labial palp at least 3-segmented. Pronotum much shorter than mesoscutum, with slight anterior collar (flange) and posteroventral angle rounded; mesoscutum transverse; notaulus distinct, complete; tegula small, convex; metapostnotum apparently about as long as metanotum; propodeum long, weakly constricted apically; meso-metapleural suture straight. Hind wing with two closed cells, vein C present only basally, anal (vannal/plical) lobe less than half length of wing. Tibiae without spines or strong setae; tibial spurs 1–2–2; basitarsomeres much longer than other tarsomeres; arolia large; claws simple. Metasoma ovoid, sessile basally, apical tergum apparently simple, seventh sternum reduced, hypopygium simple with convex apex.
Female. Unknown.
Holotype male (
Paratype male (
The species name, a noun in the genitive case, honours Professor Dr Alexandr Rasnitsyn, who first recognised the significance of the specimens.
(based on holotype, paratype data in parentheses where different or feature not visible in holotype):
Head: Hypognathous; about as wide as high; vertex evenly rounded. Eye ovate with convex inner margin, moderately protuberant, apparently glabrous, ommatidia distinct. Ocelli ovate, large. Occipital carina distinct. Frons and clypeus weakly convex; clypeus transverse with convex anterior/apical margin. Gena simple. Antennal sockets simple, apparently about as close to eyes as to each other, apparently close to posterior/dorsal margin of clypeus. Antennal scape about as long as wide (distinctly flattened posterolaterally and broadened towards apex), with several erect setae; pedicel (about half length of scape and of first flagellomere), with many fine long erect setae; (flagellomeres 1–4 becoming slightly longer sequentially, with many fine long erect setae). Mandible long, evenly broad and curved; two prominent short curved setae on lateral surface; apex truncate with four similar sharp teeth, apical tooth the longest. Maxillary palp at least 5-segmented; labial palp at least 3-segmented [palp bases concealed by foam but segmentation inferred from assumed points of origin].
Mesosoma ovate, about twice as long as wide/high. Pronotum forming a curved oblique ribbon anterolateral to mesoscutum, broader medially than posterolaterally; posterodorsal margin evenly concave; posterolateral margin strongly emarginate and approaching tegula dorsally; posteroventral angle broadly rounded; anteroventrally with slight collar (flange) but leaving propleura exposed anteriorly. Propleura closely associated or fused; anteriorly produced as a short neck; swollen posteriorly; posterior margin apparently almost straight but exposing small part of prosternum medially; forecoxae approximated. Mesoscutum shorter than wide, moderately convex; notaulus distinct and complete, weakly diverging anteriorly; tegula small and convex. (Mesoscutellum apparently almost as long as scutum, weakly convex.) Mesopleuron large and convex; meso-metapleural suture distinct, almost straight. Mesosternum with posteromedial margin almost straight and apparently slightly overhanging mesocoxal base; mesocoxae slightly separated. (Metanotum short and transverse, flattened, not constricted medially. Metapostnotum apparently as long as metanotum, slightly depressed.) Metasternum apparently somewhat depressed. Metacoxae slightly separated. Propodeum slightly longer than mesoscutum, weakly convex but more strongly so posteriorly although without any defined posterior declivity; incision between mesosoma and metasoma weak.
Forewing broad, about 2.2 × as long as wide, about 1.7 (1.8) × as long as mesosoma, with seven closed cells, veins approaching but not reaching margin. Costal cell well developed, broad. Pterostigma large, about 0.19 × as long and 0.17 (0.20) × as wide as wing, entirely sclerotised. Marginal cell about 2.38 (2.17) × as long as wide, 1.41 (1.52) × as long as pterostigma, apex acute. First submarginal cell about 2.08 (2.61) × as long as wide, 0.81 (0.80) × as long as marginal cell. Second submarginal cell almost as large as pterostigma, broadly sessile anteriorly, about 0.64 (0.57) × as long as first submarginal cell. Veins tubular except for nebulous free apical sections of M, Cu and A. No trace of any accessory vein(s) in apical membrane. Prestigmal vein (Sc+R) scarcely swollen, about 1.33 (1.37) × as long as vein 1Rs. Crossvein cu-a distinctly postfurcal, about 1.33 (1.75) × as long as 1Cu. Vein Cu2 absent, first subdiscal cell broadly open apically.
Hind wing about 0.8 × as long as forewing. (Basal and subbasal cells closed by tubular veins; costal cell open anteriorly, vein C present only basally. Veins tubular except for nebulous free apical sections of Rs, M, Cu and A. A few basal hamuli present in a cluster; about five apical hamuli. Crossvein rs-m long, about 2.67 × as long as 1Rs. Vein 1M very short, about 0.13 × as long as 2M+Cu. Crossvein cu-a distantly antefurcal, 2M+Cu about 0.57 × as long as 1M+Cu. Anal (vannal/plical) lobe apically delimited by moderate incision, lobe about 1.19 × as long as submedian cell, about 0.4 × as long as wing.) Jugal lobe absent.
Legs well developed, moderate in size; trochanters well developed and cylindrical; no trochantelli; tibiae without any spines or strong setae; basitarsomeres long, about as long as next three tarsomeres combined; all arolia large and flattened; claws simple ventrally. Foreleg with coxa subglobose, trochanter inserted apically; femur slightly swollen, with inner/anterior surface flattened; tibia with simple, weakly curved, bladelike calcar subapically. Mid- and hind legs with coxae globose, hind coxa somewhat larger than mid-coxa; tibiae each with two straight simple apical spurs, inner spur somewhat longer than outer.
Metasoma elongate oval, about 2.6 (indeterminable in paratype) × as long as wide/high; terga subequal in length. First tergum broad, weakly contracted toward base, profile evenly merging with second. First sternum apparently simple and evenly overlapping second. Seventh tergum apparently simple with apical margin convex. Seventh sternum apparently reduced and mostly concealed. Hypopygium simple, weakly convex, with narrowly rounded apical margin. Cercus apparently present, cylindrical. Genitalia with paramere apparently broadly rounded apically.
New Jersey amber containing specimens of
The cladograms resulting from the analyses using an aculeate outgroup (illustrated using
Data matrix for analyses of relationships of members of
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010100000[012] | 000021000[01] | 001[01]000202 | 000[01]1000[03]0 | 001[01]21[01]00[12] | [45]02?00010[12] | 0[34]00000100 | 002001000[02] | 0000000000 |
* |
1100010012 | 0000[01]01000 | 0010000202 | 0011100030 | 1000201001 | 000000000[01] | 0301000100 | 0000000000 | 0000022200 |
* |
0001100100 | 0000000001 | 0010100202 | 0011101033 | 5021011001 | 200-001103 | 1402100000 | 00000[01]0000 | 1000001020 |
* |
0211000100 | 0000100101 | 1100000202 | 0011211033 | 0011011001 | 620-000103 | 1412100100 | 0001000000 | 1000001020 |
|
0110001112 | 0001200000 | 1002100100 | 1002100030 | 3300000001 | 0000000000 | 0400010000 | 0001020000 | 0000121001 |
|
111001111[02] | 0001100000 | 1002100100 | 0001200012 | 3112000010 | 100-001001 | 1401000100 | 0011020000 | 0000002301 |
|
[01]110000110 | 0001200001 | 0000100011 | 1011120043 | 221[12]0101[01]1 | 0000000000 | [01][04]01000[01]00 | 0[01]2[23]0[12]0004 | 010002[12]021 |
|
0110000100 | 0001200001 | 0000100011 | 2011121043 | 2211010111 | 300-110100 | 1101010100 | 0023000004 | 0000020221 |
|
1??0000?02 | 20100??100 | 1?0?1?0??? | 00001?001? | ????1?0010 | 6100000011 | 1401010110 | 1023000000 | 0??001000? |
|
0001100111 | 2131101101 | 1100100000 | 0000100111 | 4412211000 | 6101010011 | 0001010100 | 002201[01][01]03 | 0100010000 |
|
0000011102 | 0001112220 | 1001111200 | 1000100120 | 3422211000 | 6101111010 | 1001011110 | 1123020000 | 011211100? |
|
0[12]10100111 | 2011101101 | 1100100000 | 0000100102 | 44[12][12]211000 | 6102110011 | 1001010110 | 1022011003 | 0101011000 |
|
0000001202 | 2001213200 | 0001111200 | 11001?1132 | ?411211010 | 620-011011 | 1301010110 | 0013220000 | 010001010? |
|
0000011102 | 2121113220 | 1001111200 | 1000100120 | 3412211010 | 6101011011 | 1001011110 | 1122020010 | 010111020? |
|
0000011102 | 0001112220 | 1001121200 | 10000?0120 | ?411211000 | 6101011012 | 0001011110 | 1122020000 | 010111020? |
|
0000011102 | 0001113210 | 1001111200 | 10001?0120 | ?411211000 | 6101011011 | 0001011110 | 1122020000 | 010211100? |
|
1110000100 | 2000000100 | 0020001000 | 1000101020 | 3301000011 | 620-001012 | 2222110111 | 0002000000 | 0100021020 |
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1??0000100 | 0000[02]00100 | 00200010?? | [12]0001?10?? | ?322000111 | 65[01]-00101? | 2222110111 | 0002100000 | 0100010020 |
|
1110000100 | 1000000100 | 0020001000 | 1000101020 | 3312000010 | 620-001012 | 2222110111 | 0012100000 | 0100021020 |
|
1130000122 | 20[01]0001000 | 0000100010 | 0011011032 | 4122010111 | 651-000012 | 2522110100 | 0021000004 | 0100020210 |
|
111000012[02] | 0000001000 | 0000100000 | 1011011032 | 4122010111 | 651-001111 | 2522110100 | 0021020004 | 0100010010 |
|
1[01][012]0000120 | [02]0[01]0[02]01000 | 0000100000 | 101101[01]032 | 4122010111 | 651-001111 | 2522110100 | 0021020004 | 0100020210 |
|
1000000100 | 0000111000 | 1010001000 | 0011110133 | 3122001111 | 651-001111 | 2211110100 | 0012000004 | 0100022020 |
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1110001100 | 0000221000 | 1010000010 | 0011210[01]33 | 4122001011 | 651-001011 | 1222110100 | 0013000003 | 0100022020 |
|
1110001100 | 0001011001 | 1010000000 | 00111?0033 | ?222001011 | 65[01]-001110 | 1322110100 | 0013000004 | 0100022??0 |
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111000?100 | 0001211001 | 1010000000 | 00111?0033 | ?222001011 | 651-001110 | 1222110100 | 0013000001 | 0?00022??0 |
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111000110[02] | 0001244001 | 1010001000 | 00111?0033 | ?[02]22001012 | 651-000112 | 1322110110 | 0113010004 | 0100022??0 |
|
1110001100 | 0001211001 | 1010000000 | 0011[12]10[01]33 | 4[02]12001011 | 651-00111[013] | 13[12][12]110110 | 0013000004 | 0100022020 |
|
0220001020 | 1001111100 | 1000100010 | 0011100022 | 5121110012 | 630-000012 | 2223110100 | 1022010002 | 0100010010 |
|
0220001020 | 1001132100 | 1000110010 | 0011100012 | 5121110012 | 630-000012 | 2223110100 | 1022010002 | 0100010010 |
|
1220001310 | 10001[02][12]201 | 0010000000 | 0001100132 | 5022100010 | 650-001112 | 1423110110 | 1022100002 | 0100022020 |
|
1[12][12]0000310 | 1000101[12]01 | 0000100000 | 0011100132 | 5022[01]00010 | 650-001112 | 2523110100 | 1022100002 | 0100022020 |
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[01]1[12]1000300 | [01]0000[123][12]001 | 0000120012 | 0011101[01]32 | 5[23]12100111 | 6[45][01]-00[01][01]1[012] | 1423110110 | 0022000002 | 01000[12]1020 |
*Non-aculeate outgroup taxa excluded from some analyses (see text)
To try to address the second of the above concerns, analyses were done using four non-Aculeata as outgroup. The resulting cladograms (
At first sight, consideration of all of the above results, involving not only the placement of
Preferred cladogram of families of
Apart from the above results, the variety of analyses performed has shown that the use only of an outgroup which is sister to the ingroup, and which may have many characters with relatively more-derived states than the ingroup, may produce misleading or ambiguous results (
Whether
I am most grateful to Alex Rasnitsyn for bringing the New Jersey fossils to my attention; it is with the greatest pleasure that I dedicate this paper to him in commemoration of his 75th birthday. I am also grateful to David Grimaldi (American Museum of Natural History, New York) for lending the specimens to me and for comments on some characters; to Arturo Roig-Alsina (Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires) for the donation of specimens of
All specimens are in D.J. Brothers’ collection (to be deposited in Iziko South African Museum, Cape Town (SAM), in due course) unless otherwise stated.
1. Compound eye, inner margin: sinuate = 0, convex = 1;
2. Compound eye, pores and setae: no pores or setae = 0, scattered pores and/or setae = 1, dense pores and/or setae = 2;
3. Compound eye, setae: absent = 0, minute = 1, short = 2, long = 3;
4. Antennal socket, distance from epistomal suture: less than socket width = 0, more than 1.5 × socket width = 1;
5. Clypeus, shape: transverse, without prominent median lobe = 0; with long median lobe narrower than intermandibular area = 1;
6. Clypeus, form: platelike, apical margin not deflexed = 0, convex, thickened, apical margin deflexed medially = 1;
7. Occipital carina: present = 0, absent = 1;
8. Antenna, antennomere number: more than fourteen = 0, thirteen = 1, twelve = 2, ten = 3;
9. Antenna, radicle-scape axis and insertion: axis nearly straight, simple annular constriction = 0, axis angled, simple annular constriction = 1, axis angled, radicle under flangelike expansion of scape = 2;
10. Antenna, scape form: simple, more or less cylindrical = 0, basally expanded ventromesally = 1, apically expanded ventromesally = 2 (NON-ADDITIVE);
11. Antenna, flagellomere setae orientation: inconspicuous and decumbent = 0, conspicuous and semi-decumbent, some erect = 1, conspicuous and erect = 2;
12. Antenna, flagellomere setae distribution: evenly developed, scattered = 0, better developed ventrally, in irregular transverse rows = 1;
13. Antenna, flagellomere setae length: much less than flagellomere width = 0, about 0.5–1 × flagellomere width = 1, about 1.5–2 × flagellomere width = 2, about 3–4 × flagellomere width = 3;
14. Mandible, form: apical margin truncate (chewing type) = 0, apical margin strongly tapering (cutting type) = 1;
15. Mandible, apical teeth: four or more = 0, three = 1, two or fewer = 2;
16. Maxillary palp, segments: six = 0, five = 1, four = 2, three = 3, one = 4;
17. Labial palp, segments: four = 0, three = 1, two = 2, one = 3, absent = 4;
18. Pronotum, anterior collar (flange): present, covering propleura = 0, present but reduced and exposing propleura = 1, extremely reduced to slight ridge, effectively absent = 2;
19. Pronotum, posterolateral lobe: simple = 0, with preapical vertical blunt carina, posteriorly depressed = 1, with preapical vertical lamella, posteriorly concave = 2;
20. Pronotum, posteroventral angle: rounded = 0, narrowly acute = 1;
21. Pronotum, posteroventral margin: straightish = 0, strongly concave = 1;
22. Propleura, dorsally: separated by membranous region = 0, closely approximated although not fused = 1;
23. Propleura, posterior margin: almost straight, propleura almost entirely contiguous mesad = 0, indented on medial halves, propleura partially separated = 1, mostly indented, propleura almost entirely separated = 2;
24. Proepimeron: clearly distinguishable at outer angle and along posterior margin = 0, reduced, discernible only at outer angle = 1, indistinguishable = 2;
25. Prosternum, form: mostly in a single plane = 0, distinctly biplanar, mostly depressed posteriorly = 1;
26. Prosternum, ventral view: well developed with apophyseal pit(s) = 0, visible as triangular sclerite without apophyseal pit = 1, reduced, scarcely visible = 2;
27. Procoxae, contiguity: contiguous basally = 0, well separated basally = 1;
28. Prepectus, form: very well developed, long (nearing midline) and broad = 0, well developed, short (far from midline) and broad = 1, reduced, short (far from midline) and narrow = 2;
29. Prepectus, midventrally: halves divided, free from each other = 0, halves fused midventrally = 1;
30. Prepectus, fusion: entirely articulating or surrounded by membrane = 0, fused with mesopleuron = 1, fused with pronotum = 2 (NON-ADDITIVE);
31. Notauli: diverging anteriorly = 0, parallel = 1, absent = 2 (NON-ADDITIVE);
32. Scutellum: simple = 0, produced posterodorsally as sharp-edged flange = 1;
33. Mesepimeron, development: well developed, flangelike, overlapping metepisternum = 0, much reduced, abutting metepisternum = 1;
34. Mesosternum, form posteriorly: smoothly truncate = 0, mesad with short transverse carina or weak tooth = 1, mesad with lamella projecting over mesocoxal base = 2;
35. Mesocoxae, contiguity: broadly separated = 0, slightly separated = 1, contiguous = 2;
36. Mesosoma, second dorsal phragma: strongly oblique, muscles 2ph-3ph anterior = 0, scarcely oblique posteriorly, muscles 2ph-3ph anterior = 1, scarcely oblique posteriorly, muscles 2ph-3ph posterior = 2 (NON-ADDITIVE);
37. Metanotum, shape: mesally about as long as laterally or longer = 0, mesally half as long as laterally or shorter = 1;
38. Metanotum, lateral length: less than half medial length of scutellum = 0, more than two-thirds medial length of scutellum = 1;
39. Metapostnotum, development: distinctly lengthened laterally = 0, evenly long throughout = 1, distinctly shortened laterally = 2, shortened throughout = 3, greatly expanded posteromedially = 4 (NON-ADDITIVE);
40. Metapostnotum, hind margin: entirely distinct = 0, distinct medially, indistinct laterally = 1, obliterated medially, indistinct laterally = 2, entirely obliterated = 3 (NON-ADDITIVE);
41. Mesosoma, third dorsal phragma: forming distinct even flange = 0, forming narrow even flange = 1, forming narrow even flange laterally only = 2, medially reduced but distinguishable = 3, absent medially = 4, entirely absent = 5 (NON-ADDITIVE);
42. Metapleuron, endophragmal pit and sulcus: pit posteriorly placed, sulcus angled or rounded ventral to pit = 0, pit posteriorly placed, sulcus acutely extended ventral to pit = 1, pit posteriorly placed, sulcus entirely dorsal to pit and curved = 2, pit anteriorly placed, sulcus entirely dorsal to pit and straight = 3, pit anteriorly placed, sulcus acutely extended ventral to pit = 4 (NON-ADDITIVE);
43. Metathoracic-propodeal pleural suture, dorsally: distinct and complete = 0, reduced but partly discernible = 1, obliterated = 2;
44. Metathoracic-propodeal pleural suture, ventrally: distinct and complete = 0, reduced but partly discernible = 1, obliterated = 2;
45. Metasternum, form anteromedially: not depressed = 0, weakly depressed = 1, entirely depressed = 2;
46. Metacoxal cavities: open = 0, closed = 1;
47. Propodeum, length: longer than high = 0, shorter than high = 1;
48. Propodeum, declivity: imperceptibly merging with disk, not identifiable = 0, distinct from disk = 1;
49. Forewing, tubular/nebulous veins: reaching apical margin = 0, ending before apical margin = 1;
50. Forewing, pterostigma: large and prominent = 0, medium to small but distinct = 1, very small, indistinct or absent = 2;
51. Forewing, closed cells and cell 2Cu (variable 1): ten (C, R, 1Cu, 1R1, 1M, 2Cu, 2R1, 1Rs, 2M, 2Rs) = 0, seven (C, R, 1Cu, 1R1+1Rs, 1M, 2Cu, 2R1) = 1, seven (C, R, 1Cu, 1R1, 1M, 2Cu, 2R1)= 2, six (C, R, 1Cu, 1R1, 2Cu, 2R1) = 3, seven (R, 1Cu, 1R1+1M+1Rs, 2Cu, 2R1, 2Rs, 2M) = 4, six (R, 1Cu, 1R1+1M+1Rs, 2Cu, 2R1, 2M) = 5, seven or fewer (cells 2Cu, 2M and 2Rs open or lost) = 6 (NON-ADDITIVE);
52. Forewing, closed cells and cell 2Cu (variable 2): six or more (2Cu present and closed) = 0, seven (C, R, 1Cu, 1R1, 1M, 2R1, 1Rs) = 1, six (C, R, 1Cu, 1R1, 1M, 2R1) = 2, five (C, R, 1Cu, 1R1, 2R1) = 3, four (C, R, 1Cu, 1M) = 4, three (C, R, 1Cu) = 5;
53. Forewing, costal cell: broad, wider than thickness of bounding veins = 0, narrow, as wide as thickness of bounding veins or less = 1, eliminated, veins C and Sc+R+RS fused = 2;
54. Forewing, 2nd submarginal cell (1Rs) shape (absent coded “-”): anteriorly broadly sessile = 0, shortened and anteriorly briefly sessile to briefly petiolate = 1, much reduced and anteriorly strongly petiolate = 2;
55. Forewing, marginal cell anterior margin: more than 0.7 × pterostigma width = 0, less than 0.5 × pterostigma width = 1;
56. Forewing, apical accessory vein (“RS2”): absent, not even spectral = 0, present, nebulous or spectral = 1;
57. Forewing, prestigma (vein Sc+R) form: narrow, apically less than 1.5 × width of 1Sc+R = 0, broad, apically more than 1.8 × width of 1Sc+R = 1;
58. Forewing, prestigma (vein Sc+R) length: at least as long as 1RS = 0, less than 0.5 X length of 1RS = 1;
59. Forewing, vein Cu2: present = 0, absent = 1;
60. Hind wing, basal hamuli: several, dispersed along costal margin = 0, few, concentrated into cluster = 1, one = 2, absent = 3;
61. Hind wing, tubular/nebulous veins: reaching apical margin = 0, into apical half of wing but not reaching apical margin = 1, restricted to basal half of membrane = 2;
62. Hind wing, veins C and Sc+R: both long and separated, cell C closed = 0, both long and fused = 1, C short, Sc+R absent except at base = 2, C short but distinct, Sc+R long = 3, C absent except at base, Sc+R long = 4, C absent except at base, Sc+R short = 5 (NON-ADDITIVE);
63. Hind wing, vein M+Cu: well developed, tubular = 0, distinguishable but nebulous = 1, absent = 2;
64. Hind wing, anal veins: A1 well developed, A2 present = 0, A1 well developed (more than half length anal lobe), A2 absent = 1, A1 short (less than half length anal lobe), A2 absent = 2, A1 minute, A2 absent = 3;
65. Hind wing, veins rs-m and cu-a: both present = 0, both absent = 1;
66. Hind wing, incised anal (vannal/plical) lobe: absent, at most indicated by slight notch = 0, present, distinct acute incision = 1;
67. Hind wing, anal (vannal/plical) lobe length: short, less than 0.5 × length of wing = 0, long, more than 0.6 × length of wing = 1;
68. Hind wing, jugal lobe: present, delimited apically by incision = 0, absent = 1;
69. Tarsal claws, form ventrally: toothed = 0, simple = 1;
70. Protrochanter, basal insertion: apical on coxa = 0, basolateral on coxa = 1;
71. Foretarsus, arolium: narrower than tarsal apex and shorter than claws = 0, at least as broad as tarsal apex and at least as long as claws = 1;
72. Meso- and metatarsi, arolia: well developed = 0, vestigial = 1;
73. Mesocoxa, subdivision and insertion: large basi- and disticoxites, cavities large = 0, reduced basicoxite and large disticoxite, cavities moderate = 1, much-reduced basicoxite and large disticoxite, cavities small = 2;
74. Mesotrochantellus: distinctly present = 0, reduced but discernible as complete ring = 1, much reduced, almost indiscernable, ventrally only = 2, absent = 3;
75. Mesotibia, spurs: two = 0, one = 1, none = 2;
76. Mesotibia, scattered spines: absent = 0, weak = 1, moderately strong = 2;
77. Metacoxa, specialised area of setae on ventral surface: absent = 0, present = 1;
78. Metatrochanter, specialised area of setae on ventral surface: absent = 0, present = 1;
79. Metafemur, apex: simple = 0, with toothlike projection on each side at tibial articulation = 1;
80. Metatibia, inner apical spur: simple, similar to outer spur = 0, forming calcar with weak simple dorsal carina = 1, forming calcar with dorsal setose carina = 2, forming calcar with dorsal setal fringe only = 3, forming calcar with fine dorsal pectination = 4 (NON-ADDITIVE);
81. Metasoma-propodeum attachment: ventral, between hind coxae = 0, dorsal, distant from hind coxae = 1;
82. Metasoma, T1 articulation with S1: overlapping and articulating with base of S1 = 0, narrowed and fused with base of S1 dorsolaterally = 1;
83. Metasoma, T7 surface: simple, ecarinate = 0, longitudinally carinate = 1;
84. Metasoma, T7 apical flange: absent = 0, present, narrow = 1, present, broad, 0.33 × as long as tergum = 2;
85. Metasoma, S1: ecarinate = 0, with strong median longitudinal carina = 1;
86. Metasoma, S7: well developed and exposed = 0, reduced and partly exposed = 1, much reduced and concealed = 2;
87. Metasoma, hypopygium (S8) concealment: completely exposed or almost so = 0, partially concealed = 1, completely concealed or almost so = 2;
88. Metasoma, hypopygium (S8) shape: simple with blunt to rounded apex = 0, triangular with pointed apex = 1, truncate with weakly emarginate apex = 2, peglike = 3 (NON-ADDITIVE);
89. Metasoma, cercus: present, well developed, cylindrical = 0, present but much reduced, cylindrical = 1, absent or vestigial flattened setose disk = 2;
90. Sexual dimorphism, antennomere number: absent = 0, present (♂ 13, ♀ 12) = 1.