Melaenini

Ball, George E. & Shpeley, Danny, 2005, Taxonomic review of the Tribe Melaenini (Coleoptera: Carabidae), with observations on morphological, ecological and chorological evolution, Zootaxa 1099 (1), pp. 1-120 : 1-120

Tribe Melaenini

Scaritides (in part) Dejean 1831: 471, 478–483. Ditomides (in part) Lacordaire 1854: 166–167. Coscinides Chaudoir 1876: 115–118. Nomiini (in part) Horn 1881: 130–131. Cosciniinae Bates 1892: 296.—Bedel, 1895: 15.—1897: 109. Granigerini Bedel 1900: 24.—Sloane 1923: 244–247.—Andrewes 1930: XI.—Burgeon 1935: 192. Granigerina Iakobson 1905 200. Siagonini (in part) Peringuey 1908: 276.—Erwin 1979: 592.—1984: 381. Ditomini (in part) Peringuey 1926: 607. Cymbionotini Andrewes 1933: 3 .—1935: 18.—van Emden 1936: 46.—Burgeon 1937:

402.—Kryzhanovskij 1976 (cited by Ball 1979: 93).—Erwin and Sims 1984: 355.—Erwin

1985: 468, 469.—Madge 1989: 462.—Erwin 1991: 30.—Bousquet and Larochelle 1993:

29.—Kryzhanovskij et al. 1995: 61.—Liebherr and Will 1998: 137, 150.—Lorenz 1998b: 149. Melaenini Csiki 1933: 1650 .—Andrewes 1935: 26.—van Emden 1936: 46.—Erwin 1979:

590.1985: 468–469.—Erwin and Sims 1984: 355.—Erwin 1991: 50.—Bousquet and Laro­

chelle 1993: 29.—Liebherr and Will 1998: 150.—Roig­Juñent 1998: 352, Fig. 10 .—2000: 2,

13, Figs. 14 and 15 , respectively. Melaeninae Alluaud 1934: 30 .—Basilewsky 1953: 14.—Madge 1989: 465.—Lorenz 1998b: 149. Melaenitae (subfamily) Jeannel 1941: 291–292. Cymbionotidae Jeannel 1946: 97.—Antoine 1955: 64. Cymbionotinae Basilewsky 1953: 13. Melaenitae (supertribe) Erwin 1984: 374.1985: 468, 469.—Erwin and Sims 1984: 355.—Erwin

1991: 30.—Liebherr and Will 1998: 137. Melaenidae Deuve 1993: 146 . Trechinae (in part) Lawrence and Newton 1995: 812.

Notes about synonymy. The suprageneric names Scaritides (Dejean 1831) and Ditomides (Lacordaire 1854), with which the melaenines were associated in the first part of the 19 th Century, referred to groups now recognized as paraphyletic or polyphyletic. These taxa were systematically disassembled until each comprised only the genera presently included in the Scaritini and Ditomina , respectively.

The names Coscinides and Granigerini (and derivatives depending upon formally accepted rank­designating suffixes) were invalid as applied to melaenines: Coscinides, because Coscinia Dejean 1831 was a junior homonym of Coscinia Hübner 1822 ; and the type species of Graniger Motschulsky 1864 ( G. algirinus Motschulsky 1864 ) proved to be a ditomine (tribe Harpalini ) (see International Code of Zoological Nomenclature, 1999, page 46, article 39). Two names were proposed independently, in the same year, for the melaenines: Cymbionotini (Andrewes 1933) , with type genus Cymbionotum Baudi di Selve the valid generic name; and Melaenini (Csiki 1933) , with type genus the name Melaenus Dejean, 1831 . Both Andrewes (1935) and Csiki (1933) concluded that Cymbionotum and Melaenus were tribally distinct from one another, an opinion accepted by most subsequent authors, although Erwin (1979 and 1984), with the erroneous oral advice and encouragement of G. E. Ball, treated Cymbionotini as a junior synonym of Siagonini .

Names of the suprageneric taxa are not problematic, so long as each genus is included in its own tribe. But what is the tribal name to be, with both Cymbionotum and Melaenus in the same tribe? Both tribal names were proposed in the same year (1933), but in different months: Melaenini in May and Cymbionotini in June. Therefore Melaenini has priority.

Recognition. Melaenine adults are recognized by the following combination of character states: overall size small to moderate (length ca. 4–10 mm); fronto­vertex of head with deep paraocular longitudinal groove (cf. Figs. 11A–C ) each side; mandibular scrobes each with one to several setae; submentum and gula fused (i.e., without submental­gular suture­ Figs. 7C , 13A ); prothoracic coxal cavities closed by insertion of apex of proepimeron into lateral socket of prosternum; mesothoracic coxal cavities conjunct; metepisternum and metepimeron fused (i.e., metapleuron without pleural suture); elytron with distinct plica posteriolaterally; front tibia anisochaete; and middle tibia with distal lateral setal comb. Perhaps the most distinctive feature, but one that is seen only by dissection of females, is the very long spermathecal diverticulum ( Fig. 15E ). This alone would seem sufficient to establish monophly of the Melaenini . Larvae are unknown (Grebennikov 2001: 50).

Description. Adult beetles ( Figs. 1A, 1B ) exhibiting the recognition features of Melaenini , and following. Color of body black to testaceous, elytra concolorous or bicolored. Body form terete to depressed. Antennae and mouthparts rufotestaceous, concolorous, to bicolored, proximal antennomeres black, distal antennomeres rufous, to all antennomeres black. Legs testaceous to black.

Microsculpture. Dorsal surface generally with isodiametric mesh pattern, microlines shallow, or microlines and sculpticells confined to postocular transverse impression, or scutellum; or microlines absent, surface without evident mesh pattern; or mesh pattern transverse ( Figs. 5F–G ), microlines shallow; some appendages with fimbriate sculpticells (see generic descriptions, for details).

Luster. Body surface dull, shiny, or iridescent.

Macrosculpture. Surface generally moderately densely punctate, or smooth with only elytral striae punctate.

Vestiture. Body surface glabrous (except fixed setae), or with dense to very sparse covering of testaceous setae ( Figs. 2 , 3 ).

Fixed setae (body sclerites, mouthparts, elytra). Head capsule with one or two pairs of supraorbital setae; with or without row of long setae posteriad eyes, and curved forward over eye surface. Mouthparts: labrum dorsally ( Figs. 6J , 12I ) with row of six apical setae (as), near distal margin; mandible with single seta or with several setae in scrobe ( Figs. 6A–D , ss, and Figs. 12A–D , pss, sss); maxillary stipes with two setae ( Figs. 7A–B , 12K–L , dls, vls), palpifer with single seta ( Figs. 7A–B , 12K–L , pfs); labial submentum with four or more setae ( Figs. 7C , 13A , sms, psms, ssms); mentum with pair of paramedial setae ( Figs. 7C , 13A , ms); glossal sclerite distally quadrisetose or bisetose ( Figs. 7C , 13A , gs); labial palpomere 2 bisetose (i.e., with two long setae on anterior margin, near apex). Pronotum with single pair or two pairs of lateral setae. Elytron dorsally with or without parascutellar seta, without discal setae, laterally about 20 setae in continuous umbilicate series. Abdominal sterna IV–VI each with pair of ambulatory setae or asetose; sternum VII with ( Cymbionotum , both sexes, and Melaenus females) or without pair of setae ( Melaenus males) near posterior margin.

Head. Moderately to distinctly wide, dorsal surface moderately to slightly convex, with or without distinct transverse impression posteriad eyes; frontoclypeal suture not evident, frontal impressions hardly indicated, or broad and shallow; frontovertex each side laterad eye with distinct longitudinal sulcus extended posteriorly, to transverse groove (if present); also sharp supraorbital ridge each side extended sinuously anteriorly to mandibular fossa, and here joined to apex of fronto­vertical ridge; posteriolaterally, genae rather broad, broader than antennomere 1.

Eyes ( Fig. 1A–B , 11A–C ). Rather small, convex.

Antennae. Filiform ( Fig. 1A ), rather long, extended posteriorly beyond basal third of elytra, or nearly moniliform ( Fig. 1B ), shorter, extended to about basal third of elytra. Antennomeres variously proportioned, antennomeres 5–10 symmetric, slightly asymmetric, or markedly asymmetric (for details, see descriptions of genera).

Mouthparts. ( Figs. 6A–K , 7A–D , 12A–L , 13A–C ) Labrum ( Figs. 6J–K , 12I–J ) transverse, distal margin very shallowly to moderately concave, or distinctly notched medially. Ventrally, epipharynx as in Figs. 6K and 12J : pedium (ped) triangular, either approximately equilateral or isosceles, with broad base along distal margin; parapedial projection (pp) acute or obtuse; lateral parapedial setae (lps) four to seven; extra parapedial setae (eps) either mediad or laterad parapedial ridge (pr); coeloconic sensilla (cs) relatively few to many.

Mandibles ( Figs. 6A–I , 12A–H ). Trigonal, narrowed distally and in dorsal aspect curved mediad, in lateral aspect curved ventrad; terebra (t) with or without series of short, diagonal sulci (dts, vts) and ventral condylar sulci (vcs); dorsolateral surface with dorsal groove (dgs) extended from scrobe (s); basal brush (bb) evident at base of occlusal margin; ventral surface with moderately long ventral groove (vg), bearing dense row of short microtrichia (mtr); also, secondary ventral groove (svg) and medial ventral depression (vd). Left mandible, dorsal surface ( Figs. 6B , 12B ) with sharply inflexed incisor tooth (it), terebral tooth (tt) moderately prominent, retinaculum with anterior and posterior tooth (art, prt), anterior tooth evident or not, from dorsal surface, premolar (pmt) and molar (mt) tooth evident; posterior occlusal groove (pog) distinct; ventral surface ( Figs. 6F , 12F ) of occlusal margin with molar ridge (mr) and its basal and occlusal extensions (bemr, omr). Right mandible ( Figs. 6C, 6G , 12C, 12G ) similar to left mandible in most respects, but more curved and shorter.

Maxillae ( Figs. 7A–B , 12K–L ). Standard for Carabidae , with basal trianguloid cardo, rectanguloid stipes with dorsal and ventral lobes (dls, vls), dimerous galea (g1, g2), and lacinia (lc) with prominent distal tooth (lct), and occlusal margin thickly setose; palpus of four articles, (mp1­ 4), palpomere 1 shortest and palpomere 2 longest.

Labium ( Figs. 7C–D , 13A–C ). Submentum (sm) continuous with gula (not shown); mental­submental suture (m­sms) distinct; mentum (m) transverse, with pair of prominent, broad lateral lobes (ll); epilobes (e) broad, mental tooth (mt) small, broad, anterior margin distinctly notched. Prementum: ligula, ventral aspect ( Figs. 7C , 13A ), with broad glossal sclerite (gsc) and broad longitudinal keel medially, distal margin broadly rounded, pair of paraglossae (pg), each paraglossa slender, narrow lobe, attached to glossal sclerite at base, only, surface with or without extensive covering of seta­like microtrichia ( Fig. 13B , mtr); in dorsal aspect ( Figs. 7D , 13C ), glossal sclerite tripartite, basal portion (gsbp) narrow, medial (gsmp) and distal (gsap) portions variously extended; palpiger (pge) moderately large; palpus with three articles (lp1­3), palpomere 1 short, palpomeres 2 and 3 longer, subequal, palpomere 3 broader, fusiform, apex narrowed, surface more or less setose.

Prothorax. Pronotum ( Figs. 1A, 1B ) subcordate, markedly to slightly wider than long, broader at anterior than at posterior margin; anterior margin slightly concave, posterior margin almost straight, very shallowly sinuate; lateral margins anteriorly rounded, posteriorly sinuate; anteriolateral angles subacute, projected slightly anteriorly; posteriolateral angles acute, prominently projected or not; anterior margin not beaded, flat, posterior margin beaded or not, lateral margins narrowly beaded or not; transverse discal impressions not evident, medial longitudinal impression, distinct, evident medially, not extended to either anterior or posterior margins; lateral grooves broad to narrow; disc slightly convex, lateral declivity each side gradual or rather steep. Prosternum with intercoxal process short, broad posteriorly, ventral surface with shallow longitudinal groove.

Scutellum. Trianguloid, apex pointed.

Elytra ( Figs. 1A, 1B ). Extended length of abdomen. Each elytron oblong, humerus broadly or narrowly rounded, preapically narrowed to narrowly rounded sutural apex, lateral margin straight, not sinuate; dorsal surface plane for most of length, apical declivity gradually to moderately steeply sloped; basal ridge narrow, not extended to edge of scutellum, anteriorly angulate ( Fig. 1A ), or not ( Fig. 1B ); striae not evident, or very slightly impressed, intervals flat, or striae distinct throughout length, nine in number, intervals moderately convex; parascutellar stria and diagonal portion of stria 1 absent, parascutellar setigerous puncture at base of stria 1. Plica posteriolaterally distinct.

Hind wings ( Figs. 8A , 15A ). Macropterous. Oblongum cell (o) large; wedge cell (w) large or absent. Veins RP 3 + 4 and AA 3 + 4 present or absent. Vein MP 4 more than, or less than, half­length of MP 3.

Legs. Articles of about average proportions for ground­inhabiting carabid adults. Front tibia with antenna cleaner type B (Hlavac 1971: 57); upper spur unusually large. Middle tibia distally with sculpticells ( Figs. 7F , 13D , ifs, ofs) fimbriate or not. Hind coxae narrowly in contact with one another in midline. Tarsomere 5 distoventrally ( Figs. 7G , 13E ) with prominent lobe, rounded at apex (avl); unguitractor plate (up) narrow or broad; claws (tc) with ventral margins smooth, not serrate or pectinate. Front tarsi of males without adhesive vestiture.

Abdominal sclerites. Tergum VIII partially invaginated; with anterior projections (Roig Juñent 1998: 345, Table 2, and 357). Sterna III and IV evidently fused, with intersegmental suture fine and incomplete, not extended width of abdomen. Sterna IV–VI narrowly transversely sulcate, posteriorly. Sternum VII with posterior margin rounded, and beaded.

Male genitalia ( Figs. 8B–D , 15B–D, 15F , 17D–E, 17I–K, 19C–D, 19G–H, 19K, 19O, 22D–I). Phallus with base open dorsally (Roig Juñent 1998: 345, Table 2, and 357), right basal lobe absent, most of dorsal surface membranous. Endophallus with sclerite X contained wholly within phallus, large, more than half length of phallus, or much shorter than phallus; or small, basad and partially outside phallus; or absent; parameres slender, elongate, with dense row of long setae distally ( Figs. 8C–D ), or glabrous ( Figs. 15C–D ; cf. also van Emden 1936: 45, Fig. 11 ).

Ovipositor ( Figs. 7H–K , 13F–J ). Laterotergite IX ( Fig. 15E , lt) with or without anteriorly directed apophysis. Rami present, in Melaenus about 0.80 length of gonocoxite 1, in Cymbionotum ( Fig. 15E, r ) 0.50–0.80 length of gonocoxite 1. Gonocoxa dimerous, gonocoxites 1 and 2 subequal ( Fig. 13F , gc1, 2), or gonocoxite 2 much shorter than gonocoxite 1 ( Fig. 7H , gc1, 2). Gonocoxite 1 with trichoid setae (ts) and row of ensiform setae (es). Gonocoxite 2 with subapical sensory furrow (ssf) and pair of nematiform setae (ns); without furrow pegs distad setal insertions.

Female internal genitalia ( Fig. 15E ). Bursa copulatrix (bc) elongate, voluminous; spermathecal duct (spd) with origin near junction of bursa and common oviduct (co); spermathecal duct with epispermathecal sclerite extended from bursa copulatrix to spermathecal reservoir (Liebherr and Will 1998: 118, and 169, Appendix 2); spermathecal duct shorter or much longer than spermathecal reservoir (sp), latter reniform, or sausage shaped; spermathecal reservoir with diverticulum (di) long and convoluted. Following features absent (Liebherr and Will 1998: 169, Appendix 2): accessory gland; spermatheca 2 and its appended gland; helminthoid sclerite; villous canal; sclerotized extension of spermathecal duct; diverticula of spermathecal gland; and spermathecal digitiform process.

Way of life. Little is known about this topic. Evidently, most taxa occupy drier open habitats, but the group is represented also in the tropical rain forest vegetation zone (Table 17 and Fig. 24). Bedel (1897: 109) wrote that specimens of Cymbionotum semelederi occur on sandy clay soil on desert plains, and that they come to light on warm evenings. In contrast, specimens of C. negrei Perrault were found under bark of fallen trees.

Included taxa. This tribe includes only two genera: Melaenus Dejean (with two species), and Cymbionotum Baudi di Selve (with 20 species). The classification is outlined in Table 1.

Geographical distribution ( Fig. 4 ). The range of the Melaenini includes the Oriental, southern Palaearctic (principally south of 40º N. Lat.), and Afrotropical (principally north of the Equator) Regions of the Eastern Hemisphere ; and northernmost South America , in the Neotropical Region of the Western Hemisphere .

Figures

FIGURE 1. Photographs illustrating dorsal habitus: A, Melaenus piger Fabricius (SBL 7.33 mm); B, Cymbionotum fernandezi, new species (SBL 3.96 mm).

FIGURE 2. Elytra of Cymbionotum species. SEM photographs illustrating setal length. A, left elytron, dorsal aspect, of C. fernandezi, new species. B–F, portions of left elytra, dorsolateral aspect, of: B, C. pictulum Bates; C, C. semelederi Chaudoir; D, C. striatum Reitter, E, C. fernandezi, new species; and F, C. semirubricum Reitter. Scale bars: A = 1.0 mm; B–F = 100 µm.

FIGURE 3. Elytra of Cymbionotum species. SEM photographs of sections of elytral discs in anterior one third, illustrating density of elytral punctation: A, C. striatum Reitter; B, C. semelederi Chaudoir; C, C. schueppelii Dejean; D, C. pictulum Bates; E, C. semirubricum Reitter; and F, C. fernandezi, new species. Scale bars = 100 µm.

FIGURE 4. Generalized distribution of members of the Tribe Melaenini illustrated with crosshatching on a Winkel's 'Tripel' Projection map of the world.

FIGURE 5. Antenna and elytron of Melaenus piger Fabricius illustrated with SEM photographs. A, antennomeres 1–3, anterior aspect. B, antennomere 3, illustrating ridged microsculpture. C, antennomeres 6 and 7, posterior aspect, illustrating form and setation. D, antennomere 6, illustrating setae and medial longitudinal carina. E, left elytron, dorsal aspect, illustrating form and striation. F and G, left elytron, dorsal aspect, illustrating microsculpture mesh pattern, punctation and setation. Legend: A2, A3, A6 and A7, antennomeres 2, 3, 6 and 7; ac, antennal carina; mtr, microtrichial ridge; qfs, quadrifurcate seta; sp, strial puncture. Scale bars: A, C and F = 100 µm; B, D and G = 10 µm; and E = 1mm.

FIGURE 6. Structural features of Melaenus piger Fabricius illustrated with SEM photographs. A– I, mandibles: A, D, lateral aspect, left and right mandible, respectively; B, C, dorsal aspect, left and right mandible, respectively; E, H, occlusal aspect, left and right mandible, respectively; and F, G, ventral aspect, left and right mandible, respectively; I, scrobe of left mandible, microsculpture. J, K, labrum, dorsal and ventral aspect, respectively. Legend: art, anterior retinacular tooth; as, apical seta; b; basal portion of mandible; bb; basal brush; bemr, basal extension of molar ridge; cr, crepis; cs, coeloconic sensillum; dgs, dorsal groove of scrobe; dts, dorsal terebral sulcus; eps, extra parapedial seta; fs, fimbriate sculpticell; it, incisor tooth; lps, lateral parapedial seta; mps, medial parapedial seta; mr, molar ridge; mt, molar tooth; mtr, microtrichia; omr, occlusal part of molar ridge; ped, pedium; pog, posterior occlusal groove; pp, parapedial projection; pr, parapedial ridge; prt, posterior retinacular tooth; rr, retinacular ridge; s, scrobe; so, sense organ; ss, scrobal seta; svg, secondary ventral groove; t, terebra; tr, terebral ridge; tt, terebral tooth; vcs, ventral condylar sulcus; vg, ventral groove; vts, ventral terebral sulcus. Scale bars: A–H and J–K = 100 µm; I = 10 µm.

FIGURE 7. Structural features of Melaenus piger Fabricius illustrated with SEM photographs. A, B, left maxilla, dorsal and ventral aspect, respectively; C, D, labium, ventral and dorsal aspect, respectively; E, F, left middle tibia, apical portion, and basal portion of tarsomere 1, anterior aspect, E—low magnification, F—high magnification, showing microsculpture; G, middle tarsomere 5, apical portion, and claws, ventral aspect; H–K, ovipositor, left gonocoxa—H, gonocoxa 1 and 2; I– K, gonocoxite 1, apical portion and gonocoxite 2, lateral, dorsal and ventral aspects, respectively. Legend: c, cardo; cs, coeloconic sensillum; ats, anterior tibial spur; avl, apical ventral lobe of t­5; css, clamshell sensillum; dls, dorsal lobe of stipes; e, epilobe of mentum; es, ensiform seta; g1, galeomere 1; g2, galeomere 2; gc1, gonocoxite 1; gc2, gonocoxite 2; gs, glossal apical seta; gsap, glossal sclerite, apical portion; gsbp, glossal sclerite, basal portion; gsc, glossal sclerite; gsmp, glossal sclerite, medial portion; ifs, inner face sculpticell; lc, lacinia; lct, lacinial tooth; ll, lateral lobe of mentum; lp1, labial palpomere 1; lp2, labial palpomere 2; lp3, labial palpomere 3; m, mentum; mp2, maxillary palpomere 2; mp3, maxillary palpomere 3; mp4, maxillary palpomere 4; ms, paramedial seta of mentum; m­sms, mental­submental suture; mt, mental tooth; mtb, middle tibial brush; ns, nematiform seta; ofs, outer fimbriate sculpture; pf, palpifer; pfs, palpiferal seta; pg, paraglossa; pge, palpiger; pts, posterior tibial spur; sm, submentum; sms, submental seta; ssf, subapical sensory furrow; ta1, tarsomere 1; tc, tarsal claw; up, unguitractor plate; va, ventral articulation; vls, ventral lobe of stipes; vtls, ventrolateral seta. Scale bars: A–E and G–K = 100 µm; F = 10 µm.

FIGURE 8. Structural features of Melaenus piger Fabricius illustrated with line drawings. A, left hind wing with venation, dorsal aspect. B–D, phallus and endophallus, in part. Legend: wing veins and cells—AA1+2 and AA3+4, anal veins 1+2 and 3+4, respectively; MP3 and MP4, posterior median veins 3 and 4, respectively; o, oblongum cell; RA4, anterior radial vein 4; RP2 and RP3+4, posterior radial veins 2 and 3+4, respectively; w, wedge cell. Male genitalia: x, sclerite x of endophallus. Scale bars = 1.0 mm.

FIGURE 10. Geographical range of the species of Melaenus Dejean illustrated on an outline map of southwestern Asia, southern Europe, and Africa.

FIGURE 11. Heads and antennae of Cymbionotum species illustrated with SEM photographs: A–C, head, dorsal aspect of: A, C. semelederi, Fabricius; B, C. microphthalmum Chaudoir; C, C. helferi Chaudoir. D–G, left antenna of: D, C. semirubricum Reitter (antennomeres 1–3, anterior aspect); E, C. striatum Reitter (antennomeres 1–3 and base of 4); F and G, C. semirubricum and C. striatum, respectively (antennomeres 6 and 7, posterior aspect). Legend: A2, A3, A6 and A7, antennomeres 2, 3, 6 and 7; ct, clypeal tooth; poi, postocular impression. Scale bars: A–C = 1.0 mm; D–G = 100µm.

FIGURE 12. Structural features of Cymbionotum fernandezi, new species, illustrated with SEM photographs. A–H, mandibles: A, D, lateral aspect, left and right mandible, respectively; B, C, dorsal aspect, left and right mandible, respectively; E, H, occlusal aspect, left and right mandible, respectively; and F, G, ventral aspect, left and right mandible, respectively. I, J, labrum, dorsal and ventral aspect, respectively. K,L, left maxilla, ventral and dorsal aspects, respectively. Legend: aog, anterior occlusal groove; art, anterior retinacular tooth; as, apical seta; b; basal portion of mandible; bb; basal brush; bemr, basal extension of molar ridge; c, cardo; cr, crepis; cs, coeloconic sensillum; dgs, dorsal groove of scrobe; dls, dorsal lobe of stipes; eps, extra parapedial seta; g1, galeomere 1; g2, galeomere 2; it, incisor tooth; lc, lacinia; lct, lacinial tooth; lps, lateral parapedial seta; mp2, maxillary palpomere 2; mp3, maxillary palpomere 3; mp4, maxillary palpomere 4; mps, medial parapedial seta; mr, molar ridge; mt, molar tooth; mtr, microtrichia; omr, occlusal part of molar ridge; ped, pedium; pf, palpifer; pfs, palpiferal seta; pmr, premolar ridge; pmt, premolar tooth; pog, posterior occlusal groove; pp, parapedial projection; pr, parapedial ridge; prt, posterior retinacular tooth; pss, primary scrobal seta; rr, retinacular ridge; s, scrobe; sss, secondary scrobal seta; svg, secondary ventral groove; t, terebra; tr, terebral ridge; tt, terebral tooth; vd, ventral depression; vg, ventral groove; vls, ventral lobe of stipes. Scale bars = 100 µm.

FIGURE 13. Structural features of Cymbionotum fernandezi, new species, illustrated with SEM photographs. A, labium, ventral aspect; B. paraglossa, dorsal aspect; C, labium, dorsal aspect; D, middle tibia, apical portion, anterior aspect; E, middle tarsomere 5, apical portion, and claws, ventral aspect; F–J, ovipositor, left gonocoxa––F, gonocoxa 1 and 2, lateral aspect; G–I, gonocoxite 1, apical portion and gonocoxite 2, lateral, dorsal and ventral aspects, respectively; J, gonocoxite 2, apical portion, oblique aspect. Legend: cs, coeloconic sensillum; ats, anterior tibial spur; avl, apical ventral lobe of t­5; css, clamshell sensillum; dts, dorsal trichoid seta; e, epilobe of mentum; es, ensiform seta; gc1, gonocoxite 1; gc2, gonocoxite 2; gs, glossal apical seta; gsap, glossal sclerite, apical portion; gsbp, glossal sclerite, basal portion; gsc, glossal sclerite; gsmp, glossal sclerite, medial portion; ifs, inner face sculpticell; ll, lateral lobe of mentum; lp1, labial palpomere 1; lp2, labial palpomere 2; lp3, labial palpomere 3; m, mentum; ms, paramedial seta of mentum; m­sms, mental­submental suture; mt, mental tooth; mtb, middle tibial brush; mtr, microtrichia; ns, nematiform seta; pg, paraglossa; pge, palpiger; psms, primary submental seta; sm, submentum; ssf, subapical sensory furrow; ssms, secondary submental seta; tc, tarsal claw; ts, trichoid seta; up, unguitractor plate; vls; ventrolateral seta. Scale bars: A, C– F, and I = 100 µm; B and J = 10 µm.

FIGURE 14. Pronotum, dorsal aspect, of Cymbionotum species, illustrating subgeneric differences with outline drawings. A, C. (Procoscinia) fernandezi, new species. B, C. (Cymbionotum) semelederi Chaudoir. Legend: pep, proepipleuron. Scale bars = 1.0 mm.

FIGURE 15. Structural features of Cymbionotum fernandezi, new species (A–E) and C. negrei Perrault (F) illustrated with line drawings. A, left hind wing, dorsal aspect. B, male genitalia: phallus, left lateral aspect, with endophallus partially everted; C–D left and right paramere; respectively, ventral aspect. E, female genitalia, ventral aspect. F, male genitalia, left lateral aspect. Legend: wing veins and cells­­ AA1+ 2, anal vein 1 + 2; MP3 and MP4, posterior median veins 3 and 4, respectively; o, oblongum cell; RA4, anterior radial vein 4; RP2, posterior radial vein. Male genitalia: eph, endophallus; ph, phallus; x, sclerite x of endophallus. Female genitalia: bc, bursa copulatrix; co, common oviduct; di, spermathecal diverticulum; gc 1 and 2, gonocoxite 1 and 2, respectively; lt, laterotergite of abdominal segment IX; sp, spermatheca; spd, spermathecal duct. Scale bars: A and E = 1.0 mm; and B–D and F, = 0.5 mm.