Nothoodes taprobanae ( Andrewes, 1923 ), 2020

Guéorguiev, Borislav & Liang, Hongbin, 2020, Revision of the Palaearctic and Oriental representatives of Lachnocrepis LeConte and Oodes Bonelli (Coleoptera: Carabidae), with special account on Chinese species, Zootaxa 4850 (1), pp. 1-89 : 78-82

23. Nothoodes taprobanae ( Andrewes, 1923) , comb. n.

( Figs 32 A–E, Table 5)

Oodes taprobanae Andrewes, 1923: 231 (type locality: ‘ Vavuniya’ , Sri Lanka).

References. Oodes taprobanae: Andrewes 1928: 170 (distribution in Sri Lanka); Andrewes 1930: 239 (‘ Ceylon: Vavuniya, Anuradhapura.’); Andrewes 1940: 204 (identification key; distribution: ‘Ceylon’); Döbler 1975: 144 (type material deposition); Lorenz 1998: 305; Lorenz 2005: 326.

Oodes (Oodes) taprobanae: Csiki 1931: 1011 (‘Ceylon’).

Type material. Oodes taprobanae Andrewes : holotype ♂, ‘Type H.T. [white rounded label with red band, p] // Ceylon 1922.215 [w, h] // Vavuniya ... 04. [w, h] / Oodes taprobanae Andr. Type H.E.Andrewes det. [w, h/p]’ (BMNH); paratype ♀, ‘Anuradhapura Ceylon W. Horn 1899 [w, p] // COTYPUS [r, p] // Oodes taprobanae Andr. Cotype ♀ H.E.Andrewes det. [w, h/p] // DEI Müncheberg Col–07196 [light green, p]’ (SDEI).

Other material examined. None.

TME: 2 specimens. TGE: 1♀ (only spermatheca studied; gonocoxae were not examined since terminal parts of two apical gonocoxites had been destroyed).

Diagnosis. This species is readily differentiated from other congeners by its small size, less than 12 mm, granulation in marginal furrow of elytra discontinuous, and lower EL/ PL ( Table 5).

Description (based on female paratype). Habitus. Specimen small for Nothoodes (BL: 10.1 mm, BW: 4.2 mm), with subelongate and moderately convex body ( Fig. 32A ). Ratios and measurements. See Table 5. Color and luster. Dorsal surface black, without iridescence; ventral surface brown, shiny, with iridescence; antennae, palpi, tibiae and tarsi rufopiceous. Punctuation. Dorsal surface without punctuation except posterolateral parts of pronotum, which are with large, relatively shallow punctures extended laterally to posterior half; proepisternum and ventrite 6 at apex with shallow punctures; pro- and metasternum at sides, meso- and metepisternum, and ventrite 1 at base with coarse punctures; ventrites wrinkled at sides. Head. Relatively narrow, half as wide as protonum ( Table 5). Mentum tooth with apex rounded, with paramedial border situated more apically than in other congeners ( Fig. 32B ). Thorax. Pronotum with sides barely rounded toward posterior angles; maximum width close to posterior angles; laterobasal impressions distinct, moderately impressed; base slightly sinuate. Prosternum with shallow median longitudinal sulcus; prosternal process round, with border distinct subapically and very faint at apex ( Fig. 32C ). Metepisternum longer than wide (MA/ MM: 0.92), its coadunation with epipleuron short, located anteriorly, less expressed than in N. angustatus ( Fig. 32D ). Elytra. Apical sinuation distinct, but weak. Basal margin distinct, forming a small denticle at shoulder, decreasing medially, reaching level of stria 3. Granulation in marginal furrow discontinuous, interrupted in second fifth of elytron. Parascutellar striola weakly developed, punctiform; striae 1–4 less impressed than other striae. Intervals 1–7 flat to subconvex, interval 8 convex. Male genitalia. Unknown. Female genitalia. Gonocoxite not studied (see remarks above). Bursa copulatrix sub-conical (as in N. angustatus ); spermathecal gland connected at middle of seminal canal ( Fig. 32E ).

Distribution. Oriental region: Sri Lanka.

Bionomics. Nothing is known about the bionomics of this species.

Key to species of Oodes Bonelli 2, Pseudoodes gen. n., Sundaoodes gen. n., and Nothoodes gen. n. in Palaearctic and Oriental regions

1 Submentum with one pair of setiferous punctures ( Fig. 16C ) ( Oodes Bonelli )...................................... 2

- Submentum with two pairs of setiferous punctures ( Figs 2A , 27C ).............................................. 7

2 Ventral surface of mesotarsomeres 1–4 in both sexes and protarsomeres 1–4 in female with short and relatively sparce setae aligned in one row on each side of tarsomeres ( Oodes s.str.).................................................... 3

- Ventral surface of mesotarsomeres 1–4 (or 2–4) in both sexes and protarsomeres 1–4 (or 2–4) in female with long and dense pubescence aligned in two-three rows on each side of tarsomeres ( Lachnocrepis LeConte )........................... 6 Note: Similar but less developed pilosity exists also on the ventral sides of metatarsomeres 1–4 (or 3–4).

3 Metacoxal basal sulcus extending to medial fifth of coxa ( Fig. 21G ). Stria 7 in anterior third of elytron shallower than adjacent medial striae..................................................... Oodes (Oodes) gracilis A. Villa & G.B. Villa

2 The key does not include Oodes tokyoensis Habu, 1956 for the lack of material available for study. The morphology of this taxon shows great similarity with that of O. integer due to which its distinction is unreliable.

Note: For additional differences between O. helopioides and O. gracilis see Fontolan (1959: 121).

- Metacoxal basal sulcus extending to lateral third of coxa ( Figs 16G , 18B, D ). Stria 7 in anterior third of elytron as developed as adjacent medial striae.................................................................................. 4

4 Larger species, BL>12.0 mm and BW> 5 mm. PW /PA>1.94. PA/PB <0.54....... Oodes (Oodes) echigonus Habu & Baba

- Smaller species, BL <12.0 mm and BW <5 mm. PW /PA <1.94. PA/PB> 0.54...................................... 5

5 PW / PL >1.50. EL/EW ≤1.46. Granulation in elytral marginal furrow continuous. Apex of prosternal process shorter, subovate ( Fig. 18J ). Smaller species, BL <10 mm .................................... Oodes (Oodes) helopioides (Fabricius)

- PW / PL <1.50. El/EW>1.46. Granulation in elytral marginal furrow discontinuous, separated into shorter, humeral, and longer, apical group. Apex of prosternal process longer, subpointed ( Fig. 20E ). Larger species, BL> 10 mm ............................................................................................... Oodes (Oodes) integer Semenov

6 Antennae, palpomeres, tibiae and tarsomeres black. PW /HW>1.77. MA/ MM ≤0.90. Ventral side of metatarsomere 1 in both sexes glabrous at middle. Ventral side of mesotarsomere 1 in both sexes glabrous or with single setae at middle on distal third. Ventral side of protarsomere 1 in female glabrous at middle or with a few distal setae............................................................................................... Oodes (Lachnocrepis) desertus Motschulsky

- Antennae, palpomeres, tibiae and tarsomeres reddish. PW /HW <1.77. MA/ MM ≥0.90. Ventral side of metatarsomere 1 in both sexes with setae in distal half. Ventral side of mesotarsomere 1 in both sexes with long, dense setae at middle on distal two thirds. Ventral side of protarsomere 1 in female with dense setae at middle........ Oodes (Lachnocrepis) japonicus (Bates)

7 Abdominal ventrite 3 with ambulatory setae ( Fig. 27F )....................................................... 8

- Abdominal ventrite 3 without ambulatory setae ( Pseudoodes gen. n.)........................................... 13

8. Pronotum with basal setiferous punctures more or less medially remote from posterior angles ( Figs 27G , 31J ) (setae are frequently absent and then punctures are hardly visible!). Protarsomere 3 in male subtrapezoidal, with sides constricted distally ( Figs 27A , 29D ). PW / PL <1.65 ( Nothoodes gen. n.).......................................................... 9

- Pronotum without basal setiferous punctures. Protarsomere 3 in male rectangular, with sides nearly parallel. PW / PL >1.65 ( Sundaoodes gen. n.)................................................................................. 12

9 Smaller species, BL <11 mm. EL/ PL <2.50. Granulation in elytral marginal furrow discontinuous, separated into humeral and apical group............................................................. Nothoodes taprobanae (Andrewes)

- Larger species, BL> 11 mm. EL/ PL >2.50. Granulation in elytral marginal furrow continuous........................ 10

10 PW /HW>1.90. EW/ PW <1.11. Elytral striae slightly impressed notably in their anterior half ( Figs 31A, I ).......................................................................................... Nothoodes longus (Andrewes)

- PW /HW <1.85. EW/ PW >1.11. Elytral striae well impressed.................................................. 11

11 Larger species, BL> 14.5 mm. Mentum tooth with apex pointed ( Fig. 27B ). Metepisternum densely punctate ( Fig. 27E )............................................................................... Nothoodes angustatus (Lorenz)

- Smaller species, BL <14.0 mm. Mentum tooth with apex rounded to subtruncate ( Fig. 29C ). Metepisternum more sparsely punctate ( Fig. 29G )................................................................. Nothoodes bharat sp. n.

12 Apophysis of prosternal process ovate, with apex widely rounded ( Fig. 14D ). Metepisternum nearly smooth ( Fig. 14E ). PW /PА <1.90. EL/EW <1.40. EL/ PL <2.67............................................... Sundaoodes hainanensis sp. n.

- Apophysis of prosternal process rhomboid, with apex sub-pointed ( Fig. 15E ). Metepisternum coarsely punctured ( Fig. 15F ). PW /PА>1.90. EL/EW>1.40. EL/ PL >2.67.................................... Sundaoodes kalimantanensis sp. n.

13. Dorsal surface bluish. Mesotarsomeres 2–4 and metatarsomeres 3–4 in both sexes as well protarsomeres 2–4 in female with long, dense, and yellowish pubescence on ventral side............................ Pseudoodes coelestinus (Chaudoir)

- Dorsal surface dark brown to black. Meso- and metatarsomeres in both sexes as well protarsomeres 3–4 in female with short, scattered brownish pubescence on ventral side............................................................. 14

14 Larger species, BL> 15 mm ............................................................................ 15

- Smaller species, BL <15 mm ........................................................................... 16

15 PW /HW>2.10. PW /PA>1.90. Metepisternum less densely punctured; MA/ MM <0.85 ( Fig. 8D )................................................................................................. Pseudoodes ampliusculus sp. n.

- PW /HW <2.00. PW /PA <1.80. Metepisternum more densely punctured; MA/ MM about 1.00 ( Fig. 3E )....................................................................................... Pseudoodes subcoriaceus (Chaudoir)

16 MA/ MM <1.00. Winged species, with loose elytra and metathoracic flight wings more or less well developed (“ vicarius ” group)............................................................................................. 17

- MA/ MM >1.40. Wingless species, with coalesced elytra and metathoracic flight wings reduced (“ rambouseki ” group)...... .................................................................................................. 18

17 Mentum tooth with apex emarginate ( Figs 4D, E ). Apophysis of prosternal process ovate ( Fig. 4F ). MA/ MM >0.93..................................................................................... Pseudoodes vicarius (Bates)

- Mentum tooth with apex subtruncate to subrounded ( Fig. 2A ). Apophysis of prosternal process elongate ( Fig. 2B ). MA/ MM <0.93..................................................................... Pseudoodes cribristernis (Bates)

18 Pronotum narrower compared with head ( PW /HW <2.10).................................................... 19

- Pronotum wider compared with head ( PW /HW>2.10)....................................................... 20

19 Shaft of median lobe very long, narrow ( Fig. 9F ); apical lamella nearly straight ( Fig. 9G ). Pronotum wider ( PW / PL >1.63)............................................................................ Pseudoodes emeishanicus sp. n.

- Shaft of median lobe shorter and wider ( Fig. 12C ); apical lamella narrower, left-orientated ( Fig. 12D ). Pronotum narrower ( PW / PL <1.63)........................................................... Pseudoodes rambouseki (Jedlička)

20 Metepisternum longer (MA/ MM <1.58).................................................................. 21

- Metepisternum shorter (MA/ MM ≥1.60)........................................... Pseudoodes hunanensis sp. n.

21 Smaller specimens, BL <13.0 mm. Pronotum longer ( PW / PL >1.60), less pronouncedly narrowed to apex ( PW /PA <1.91) ( Figs 11A, H )................................................................. Pseudoodes leigongshanicus sp. n.

- Larger specimens, BL> 13.2 mm. Pronotum shorter ( PW / PL <1.60), more pronouncedly narrowed to apex ( PW /PA>1.91) ( Fig. 13B )....................................................................... Pseudoodes tianlinensis sp. n.

Overview of the results

In order to revise the Palaearctic and Oriental representatives of Lachnocrepis and Oodes , we studied available material of 28 valid species, two species from the first genus and 26 from the second ( Bousquet 2017; Lorenz 2019). Latest figures represent all hitherto known representatives from both genera in the biogeographical regions mentioned. Some material from species belonging to genera Holosoma and Simous were included in the study.Several new synonymies have been established, among them that of Oodes desertus with O. prolixus . The morphological analysis showed that two species of Lachnocrepis and 12 species of Oodes share important taxonomic features. To this group, eight species new to science and one previously classified in Holosoma were added. All these species are classified into the “ Oodes generic group”, to demonstrate their close relations and to delimit them from other oodines. The distribution of some character states and their polarity among species of this generic series provided evidences of monophyly of four groups considered, namely Pseudoodes , Sundaoodes , Oodes , and Nothoodes . Based on structural similarities in the female reproductive tract and basal bulb of median lobe of the aedeagus, the species of Oodes , Pseudoodes , and Sundaoodes are likely more closely related to each other than any of these are to species of Nothoodes .

In connection with this study, primary types of the 13 remaining Oriental species of Oodes were examined too, and it was established that they belong to other lineages of the tribe. Until these taxa can be revised, they are provisionally placed in “tribe Oodini , species incertae sedis ”.

On another side, three Afrotropical, 20 Australasian species, the Nearctic O. amaroides , and the only extinct member of the tribe so far known O. kachinensis Liu, 2014 ( Liu et al. 2014) seem morphologically similar to those from the genera treated in this study. For this reason, the last twenty-five species are placed in Oodes as “ Oodes species incertae sedis” (see “Checklist of taxa from the Oodes generic group”). Preliminary study suggests that O. congoensis Burgeon may be the only Afrotropical representative of the same generic series. According to the original description and the structure of the prosternal process ( Basilewsky 1953: 162), this species seems to be close to members of Nothoodes . Most probably, O. lenis Péringuey and O. palpalis Klug do not belong to the series in question. Data on external morphology obtained from scanty material at hand suggest that some New Guinean and Australian oodines presently placed in Oodes may be more closely related to species from newly described genera. A new study of O. kachinensis may clarify its systematic position because: “the exoskeletons of the insect fossils are very often artificially altered due to geological pressure” (J. Schmidt, pers. comm.).

Figures

FIGURE 2. Pseudoodes cribristernis Bates, 1892 (A–B: female specimen, China, Yunnan Prov., Fugong County, Guquan Village; C: male specimen, same locality; D–E: female specimen, Thailand, Chiang Dao). A: mentum; B: prosternal process; C: right protarsomeres; D: left gonocoxite, ventral view; E: spermathecal complex and gonocoxites, ventral view. Scale lines = 0.5 mm (Figs A–C, E); = 0.2 mm (Fig. D).

FIGURE 3. Pseudoodes subcoriaceus Chaudoir, 1882, holotype. A: habitus and labels of holotype; B: head and pronotum; C: mentum; D: prosternal process; E: right metepisternum and metacoxa (arrow shows lateral end of metacoxal basal sulcus). Scale lines = 2 mm (Fig. A); = 1 mm (Figs B–E).

FIGURE 4. Pseudoodes vicarius Bates, 1873 (A–B: male specimen, Japan, Chiba Pref., Ojyaga-ike; C: female specimen, same locality; D–G: female specimen, Japan, Ishikawa Pref., Tomurobessho). A, C: habitus; B: right protarsomeres; D: labium, with emphasis to mentum; E: labium, with emphasis to submentum (white spots show locations of submental setiferous punctures); F: prosternal process; G: right metepisternum and metacoxa (arrow shows lateral end of metacoxal basal sulcus). Scale lines = 2 mm (Figs A, C); = 1 mm (Figs B, G); = 0.5 mm (Figs D–F).

FIGURE 8. Pseudoodes ampliusculus sp. n., holotype. A: habitus and labels; B: mentum (white spots show locations of submental setiferous punctures); C: prosternal process; D: right metepisternum and metacoxa (arrow shows lateral end of metacoxal basal sulcus); E: left gonocoxite, ventral view; F: spermathecal complex and gonocoxites, ventral view. Scale lines = 2 mm (Fig. A); = 0.5 mm (Fig. B, F); = 1 mm (Figs C–D); = 0.2 mm (Fig. E).

FIGURE 9. Pseudoodes emeishanicus sp. n., holotype. A: habitus and labels; B: head and pronotum; C: mentum (white spots show locations of submental setiferous punctures); D: prosternal process; E: right metepisternum and metacoxa; F: median lobe of aedeagus, left lateral view; G: same, dorsal view. Scale lines = 2 mm (Fig. A); = 1 mm (Figs B, E); = 0.5 mm (Figs C–D, F–G).

FIGURE 11. Pseudoodes leigongshanicus sp. n. (A–G:holotype; H–J:paratype).A:habitus andlabels; B: mentum; C: prosternal process; D: right metepisternum and metacoxa; E: median lobe of aedeagus, left lateral view; F: same, anterior dorsal view; G: same, posterior dorsal view; H: habitus of paratype; I: left gonocoxite, ventral view; J: spermathecal complex and gonocoxites, ventral view. Scale lines = 2 mm (Figs A, H); = 0.5 mm (Figs B–C, E–G, J); = 1 mm (Figs D); = 0.2 mm (Fig. I).

FIGURE 12. Pseudoodes rambouseki (Jedlička, 1931) (A: lectotype of Holosoma rambouseki and its labels; B–D:topotype male specimen, China, Jinfo Shan, 29°01N 107°14E; E–H: topotype female specimen, China Jinfo Shan, N29.04043 E107.18410; I–J: topotype female specimen, SE Sichuan Jinfo Shan, 29°01N 107°14E). A–B, Е: habitus; C: median lobe of aedeagus, left lateral view; D: same, dorsal view; F: mentum; G: prosternal process; H: left metepisternum and metacoxa; I: left gonocoxite, ventral view; J: spermathecal complex and gonocoxites, ventral view. Scale lines = 2 mm (Figs A–B, E); = 1 mm (Fig. H); = 0.5 mm (Figs C–D, F–G, J); = 0.2 mm (Fig. I).

FIGURE 13. Pseudoodes tianlinensis sp. n., holotype. A: habitus and labels; B: head and pronotum; C: mentum; D: prosternal process; E: left metepisternum and metacoxa; F: median lobe of aedeagus, left lateral view; G: same, dorsal view. Scale lines = 2 mm (Fig. A); = 1 mm (Figs B, E); = 0.5 mm (Figs C–D, F–G).

FIGURE 14. Sundaoodes hainanensis sp. n. (A–H:holotype; I–J: female paratype).A: habitus and labels; B: head and pronotum; C: mentum; D: prosternal process; E: left metepisternum and metacoxa; F: median lobe of aedeagus, left lateral view; G: same, ventral view; H: same, dorsal view; I: left gonocoxite, ventral view; J: spermathecal complex and gonocoxites, ventral view. Scale lines = 2 mm (Fig. A); = 1 mm (Figs B, E); = 0.5 mm (Figs C–D, F–H, J); = 0.2 mm (Fig. I).

FIGURE 15. Sundaoodes kalimantanensis sp. n. (A, C–D, G–H: holotype; B, E–F, I–K: paratype). A: habitus and labels; B: habitus; C: head and pronotum; D: mentum (white spots show locations of submental setiferous punctures); E: prosternal process; F: left metepisternum and metacoxa; G: median lobe of aedeagus, left lateral view; H: same, anterior half, dorsal view; I: left gonocoxite, ventral view; J: spermathecal complex and gonocoxites, dorsal view; K: same, ventral view. Scale lines = 2 mm (Figs A–B); = 1 mm (Figs C, F); = 0.5 mm (Fig. D–E, G–H, J–K); = 0.2 mm (Fig. I).

FIGURE 16. Oodes echigonus Habu & Baba, 1960 (A: topotype male specimen, Japan, Niigata Pref., Katamachi; B–C:topotype female specimen, same locality; D–E: male specimen, China, imprecise locality; F–G: female specimen, China, imprecise locality). A–B, D, F: habitus; C: mentum; E: prosternal process; G: left metepisternum and metacoxa. Scale lines = 2 mm (Figs A–B, D, F); = 0.5 mm (Figs C); = 1 mm (Figs E, G).

FIGURE 18. Oodes helopioides (Fabricius, 1792) (A–B: Oodes parallelus Motschulsky, 1858, lectotype and its labels; C–D: Oodes parallelogrammus Motschulsky, 1858, lectotype and its labels; E–F: Oodes thessalonicensis Schatzmayr, 1909, lectotype and its labels; G: Oodes helopioides var. fiorii Porta, 1923, lectotype and its labels; H: male specimen, Bulgaria, Slānčev brjag; I: female specimen, Strandza Mt., Indipaskha; J: male specimen, France, les Collettes).A, C, E, G–H: habitus; B: left metacoxa; D: right metepisternum and metacoxa; F: left protarsomeres; I: mentum; J: prosternal process. Scale lines = 2 mm (Figs A, C, E, G–H); = 1 mm (Figs B, D); = 0.5 mm (Figs F, I–J).

FIGURE 20. Oodes integer Semenov, 1889, Russia, Primorski Kraj, Kommisarovka River (A–B: male specimen; C–H: female specimen).A, C: habitus B: pronotum and head; D: mentum; E: prosternal process; F: left metepisternum and metacoxa (arrow shows lateral end of metacoxal basal sulcus); G: left gonocoxite, ventral view; H: spermathecal complex and gonocoxites, ventral view. Scale lines = 2 mm (Figs A, C); = 1 mm (Figs B, D, F); = 0.5 mm (Figs E, H); = 0.2 mm (Fig. G).

FIGURE 21. Oodes gracilis A. Villa & G.B. Villa, 1833 (A–B: neotype and its labels; C–D: female specimen, Bulgaria, Pomorie-See; E–G: male specimen, Albania, Mali i Tabaroshit).A, C: habitus; B: left protarsomeres; D: pronotum and head; E: mentum; F: prosternal process; G: left metepisternum and metacoxa (arrow shows lateral end of metacoxal basal sulcus). Scale lines = 2 mm (Figs A, C); = 0.5 mm (Figs B, D, F–G); = 0.2 mm (Fig. E).

FIGURE 27. Nothoodes angustatus Lorenz, 1998 (A–F: male specimen, Nepal, Siddharthanagar; G: female specimen, Nepal, 8 km NE Nepalgunj). A, G: habitus (arrow shows position of right basal setiferous puncture of pronotum); B: labium, with emphasis to mentum; C: labium, with emphasis to submentum, seen from the rear (white spots show locations of submental setiferous punctures); D: prosternal process (arrow shows place of disappearance of bordering); E: right metepisternum and metacoxa (arrow shows lateral end of metacoxal basal sulcus); F: abdominal ventrites (III–VI—number of respective ventrites; arrow shows position of ambulatory seta of ventrite III). Scale lines = 2 mm (Figs A, G); = 0.5 mm (Figs B–D); = 1 mm (Figs E–F).

FIGURE 29. Nothoodes bharat sp. n. (A–C: holotype; D: male paratype, Dwarakapuram Village; E–G: female paratype, Dwarakapuram Village). A: habitus and labels; B: head and pronotum; C: mentum; D: right protarsomeres; E: habitus; F: prosternal process; G: left metepisternum and metacoxa (left arrow shows lateral end of metacoxal basal sulcus, right one shows setiferous puncture of metatrochanter). Scale lines = 2 mm (Figs A, E); = 1 mm (Figs B, D, G); = 0.5 mm (Figs C, F).

FIGURE 31. Nothoodes longus Andrewes, 1940 (A–H: male paratype; I–J: male specimen, India, Dibrugarh). A, I: habitus; B: mentum; C: prosternal process; D: right metepisternum and metacoxa (arrow shows lateral end of metacoxal basal sulcus); E: median lobe of aedeagus, left lateral view; F: same, anterior dorsal view; G: same, ventral view; H: same, posterior dorsal view; J: head and pronotum. Scale lines = 2 mm (Figs A, I); = 0.5 mm (Figs B–C, E–H); = 1 mm (Figs D, J).

FIGURE 32. Nothoodes taprobanae Andrewes, 1923, female paratype. A: habitus, with left antennomeres 6–11 attached to separate cardboard; B: mentum; C: prosternal process; D: left metepisternum and metacoxa (arrow shows lateral end of metacoxal basal sulcus); E: spermathecal complex and gonocoxites, ventral view. Scale lines = 2 mm (Fig. A); = 0.5 mm (Figs B–E).