Zyras, STEPHENS, 1835

Assing, Volker, 2017, On Zyras sensu strictu in the East Palaearctic and Oriental regions, with a focus on the faunas of the Himalaya, India, Sri Lanka, Thailand, and Sulawesi (Coleoptera: Staphylinidae: Aleocharinae: Lomechusini), Beiträge Zur Entomologie = Contributions to Entomology 67 (1), pp. 117-192 : 120-125

3.2 Key to the species of the subgenus Zyras of the Himalaya, India, and Sri Lanka

In total, 25 named and two undescribed species are currently known from the region including the Himalaya (exclusive of China), India, and Sri Lanka. Zyras pindarae , a species of highly variable coloration, keys out at three couplets.

1. Whole body with dense and long and sub-erect to erect pubescence (in dorsal view best visible on the posterolateral contours of the head, on the lateral margins of the elytra, and on the posterior margins of the sternites) (e.g., Figs 59, 62–64 ). Body broad and with rather large abdomen (e.g., Figs 95–97 ). Antennomere XI very short, much shorter than the combined length of antennomeres IX and X, and of broadly conical shape (e.g., Figs 12–13, 21 ). Apical lobe of paramere very short (e.g., Fig. 184 ). Zyras hirtus group. .................................................................... 2

– At least the forebody with distinctly less dense and less conspicuous pubescence. Habitus more slender. Antennomere XI mostly more elongate and not of distinctly conical shape. Apical lobe of paramere usually longer and more slender. ...................................................................................................................................................................... 6

2. Forebody black; abdomen blackish-brown to black. .................................................................................................... 3

– Pale-coloured or bicoloured species. .............................................................................................................................. 4

3. Meso- and metafemora distinctly bicoloured, basally yellow and apically infuscate; antennomere XI blackish, not distinctly paler than antennomeres IV–X ( Fig. 13 ). Tergites III–V without distinct micropunctation. Median lobe of aedeagus subapically more strongly angled in lateral view and apically less acute ( Figs 178–179 ). Distribution: North India, Nepal ( Map 7 ). ...................................................................................................................... hirsutiventris

– Meso- and metafemora uniformly dark-yellowish; apices very indistinctly darker at most; antennomere XI dark-reddish, somewhat contrasting with the blackish antennomeres IV–X ( Fig. 23 ). Tergites III–V with distinct micropunctation. Median lobe of aedeagus subapically weakly angled and apically very acute both in lateral and in ventral view ( Figs 182–183 ). Distribution: Northeast India ( Map 8 ). ....................................................... luteipes

4. Meso- and metafemora distinctly bicoloured, basally yellowish and apically infuscate. Elytra bicoloured, darkyellowish with the postero-lateral portion blackish ( Fig. 60 ). Abdomen bicoloured, with segments II–V reddish and VI–X blackish ( Fig. 101 ). Distribution: South India (Map 79). ........................................................ nilgiriensis

– Meso- and metafemora uniformly dark-yellowish to reddish; apices very indistinctly darker at most. Coloration of body different. ............................................................................................................................................................... 5

5. Antennae distinctly bicoloured, blackish with the apical three antennomeres pale yellowish ( Fig. 40 ). Head and pronotum dark-brown to blackish; elytra reddish with the postero-lateral portions infuscate ( Fig. 62 ). Abdomen bicoloured, reddish with segments V–VIII blackish ( Fig. 60 ). Antenna very long (approximately 3.0 mm) and slender ( Fig. 40 ). Median lobe of aedeagus as in Figs 174–175 . Distribution: North India, Nepal ( Map 7 ). .......... ................................................................................................................................................................................. gardneri

– Antennae blackish-brown to blackish, with usually only antennomere XI, rarely X–XI or IX–XI, slightly paler ( Fig. 21 ). Body more or less uniformly reddish to reddish-brown, with abdominal segments VI–VII slightly darker at most ( Figs 59 , 95 ). Antenna much shorter (approximately 2.0 mm) and more strongly incrassate ( Fig. 21 ). Median lobe of aedeagus as in Figs 170–171 . Distribution: South India, Sri Lanka ( Map 7 ). ....................... hirtus

6. Forebody and abdomen completely blackish. Legs bicoloured with the femora blackish-brown to black and the tibiae yellowish. ................................................................................................................................................................. 7

– Coloration different, at least the abdomen not completely blackish. ......................................................................... 9

7. Forebody with blueish hue ( Fig. 50 ). Antennae blackish with antennomeres I–II and the base of III yellowish ( Fig. 22 ). Distribution: North India ( Map 2 ). ............................................................................................ nigroaeneus

– Forebody without blueish hue. Antennae, including the basal antennomeres, uniformly blackish, or nearly so. ...................................................................................................................................................................................... 8

8. Tibiae blackish, as dark as femora. Ventral process of aedeagus basally broader in ventral view ( Figs 263–264 ). Paramere much longer than median lobe and with rather long and slender apical lobe ( Fig. 265 ). Distribution: Central Nepal ( Map 4 ). ........................................................................................................................................ morulus

– Tibiae distinctly paler than femora. Ventral process of aedeagus basally more slender in ventral view ( Figs 125–128 ). Paramere approximately as long as median lobe and with short apical lobe ( Fig. 129 ). Distribution: North India, Nepal ( Map 2 ). ..................................................................................................................................................... pindarae

9. Abdomen black with the apex (segments VII–X or VIII–X) reddish, strongly contrasting with the anterior segments; forebody black. .............................................................................................................................................. 10

– Abdomen of different coloration; forebody usually not uniformly black. .............................................................. 12

10. Segment VII black or reddish ( Figs 82–85 ). Pronotum with slightly irregular and denser punctation; elytra with dense and nearly regularly distributed punctation ( Fig. 44 ). Aedeagus as in Figs 125–129 . Distribution: North India, Nepal ( Map 2 ). .......................................................................................................................................... pindarae

– Segment VII reddish ( Fig. 79 ). Pronotum with sparse and distinctly irregular punctation; elytra with sparser (posteriorly very sparse) and more distinctly irregular punctation ( Fig. 42 ). ........................................................ 11

11. Legs uniformly pale-yellowish. Median lobe of aedeagus as in Figs 117–118 . Distribution: North India, Nepal ( Map 1 ). ................................................................................................................................................................... kraatzi

– Legs bicoloured, with the femora blackish-brown to black and the tibiae and tarsi distinctly paler, yellowish to brown. Median lobe of aedeagus as in ASSING (2016a: figures 268–269). Distribution: Pakistan, Afghanistan ( Map 1 ). .................................................................................................................................................................. iniquus

12. Pronotum bright-reddish, strongly contrasting with the blackish head. ................................................................. 13

– Pronotum brown to black. ............................................................................................................................................. 17

13. Elytra bicoloured. ............................................................................................................................................................ 14

– Elytra uniformly blackish-brown to black, sometimes with blueish hue. ............................................................... 15

14. Elytra blackish with the suture and the area near the scutellum extensively yellowish-red ( Fig. 54 ). Antennomeres VIII–X distinctly transverse ( Fig. 16 ). Male sexual characters unknown. Distribution: South India ( Map 7 ). ...... ................................................................................................................................................................................. hastatus

– Elytra blackish with the anterior portion more or less extensively yellowish-red; posterior portion of suture blackish ( Fig. 57 ). Antennomeres VIII–X oblong ( Fig. 19 ). Aedeagus as in Figs 161–163 . Distribution: South India ( Map 7 ). ................................................................................................................................................... latilobatus

15. Antennae short, 1.3–1.4 mm long, and distinctly incrassate apically; antennomere X at least twice as broad as long ( Figs 17–18 ). Pronotum approximately 1.15 times as broad as long ( Figs 55–56 ). Aedeagus as in PACE (2011: figures 39–40). Small species; length of forebody 2.0– 2.2 mm. Distribution: India, Sri Lanka ( Map 7 ). ................ .......................................................................................................................................................................... parageminus

– Antennae longer and more slender,> 1.7 mm long; antennomere X less than twice as broad as long. Pronotum more slender, or if similarly transverse, pronotum, elytra, tergite II, and anterior impressions of tergites III–V coarsely punctate. Larger species, length of forebody> 2.2 mm. ............................................................................. 16

16. Antennae blackish ( Fig. 20 ). Pronotum, elytra, abdominal tergite II, and anterior impressions of abdominal tergites III–V with coarse punctation ( Figs 58 , 99 ). Pronotum more transverse, 1.15–1.18 times as broad as long ( Fig. 58 ). Rather large and robust species; length of forebody 3.0– 3.3 mm. Aedeagus as in Figs 166–167 . Distribution: Oriental and southern East Palaearctic regions ( Map 8 ). .................................................................... alternans

– Antennae brown to blackish-brown with the apical 1–4 antennomeres yellowish ( ASSING 2016a: figures 33–35). Pronotum, elytra, and anterior abdominal tergites with fine shallow punctation ( ASSING 2016a: figures 99–100, 109, 157–158, 165). Pronotum more slender, 1.03–1.10 times as broad as long. Aedeagus: ASSING (2016a: figures 274–281). Distribution: Oriental and southern East Palaearctic regions ( Map 8 ). ........................ geminus

17. Metafemora bicoloured, yellowish with distinctly infuscate apices. ........................................................................ 18

– Metafemora of uniformly pale or dark coloration. ..................................................................................................... 19

18. At least profemora, often also mesofemora of uniformly yellowish coloration. Antennomeres IV–VII distinctly oblong ( Fig. 9 ). Eyes approximately as long as distance from posterior margin of eye to posterior constriction of head ( Figs 48–49 ). Pronotum> 1.1 times as broad as long and with straight lateral margins in posterior half ( Figs 48–49 ). Punctation of pronotum and elytra irregular and moderately coarse ( Figs 48–49 ). Abdominal tergites III–VI with few setiferous punctures at posterior margins ( Fig. 94 ). Male sternite VIII conspicuously acute posteriorly ( Fig. 147 ). Median lobe of aedeagus as in Figs 142–143 . Distribution: North India, Nepal ( Map 4 ). .. .............................................................................................................................................................................. condignus

– Apices of all femora blackish. Antennomeres VI–VII transverse ( Fig. 25 ). Eyes more than twice as long as distance from posterior margin of eye to posterior constriction of head ( Fig. 51 ). Pronotum <1.1 times as broad as long and with sinuate lateral margins in posterior half ( Fig. 51 ). Punctation of pronotum and elytra regular and fine ( Fig. 51 ). Abdominal tergites III–VI with numerous setiferous punctures at posterior margins. Male sternite VIII convex posteriorly ( Fig. 149 ). Median lobe of aedeagus as in PACE (1986: figures 54–55). Distribution: Central Nepal ( Map 3 ). ...................................................................................................................................................... morvani

19. Pronotum with uneven surface and conspicuously irregular punctation more or less grouped in clusters, these clusters at least partly situated in more or less distinct impressions ( Fig. 65 ). Male tergite VIII with four blunt projections posteriorly ( Fig. 196 ). Median lobe of aedeagus as in Figs 191–194 . Distribution: Oriental and southern East Palaearctic regions ( Map 6 ). ..................................................................................................................... proximus

– Pronotum with smooth surface without impressions (except the usual postero-median impression). Punctation of pronotum regular or less conspicuously irregular. Male tergite VIII often with a median concavity, but without four blunt projections. .................................................................................................................................................... 20

20. Male sternite VIII distinctly truncate posteriorly. Ventral process of aedeagus with long, slender, and acute apex in lateral view. Apical lobe of aedeagus long and slender. ......................................................................................... 21

– Male sternite VIII convex posteriorly. Ventral process of aedeagus with shorter, less slender, and less acute apex in lateral view. Apical lobe of aedeagus distinctly shorter. ......................................................................................... 22

21. Antennae approximately 2.0 mm long and not particularly slender, antennomeres VI–IX transverse ( Fig. 24 ). Pronotum broader, approximately 1.15 times as broad as long, lateral margins straight or weakly convex in posterior half in dorsal view; pronotal punctation moderately irregularly distributed ( Fig. 53 ). Non-setiferous punctation of the anterior impressions of tergites III–V fine ( Fig. 89 ). Median lobe of aedeagus with moderately slender and acute apex in lateral view ( Figs 156–158 ). Distribution: Northeast India ( Map 9 ). ........ longilobatus

– Antennae very long and slender, 2.4–2.6 mm long; antennomeres VI–XI oblong ( Fig. 11 ). Pronotum slender, approximately 1.1 times as broad as long, lateral margins sinuate in posterior half in dorsal view; lateral portions of disc with a cluster of punctures slightly behind middle, otherwise nearly impunctate ( Fig. 52 ). Non-setiferous punctation of anterior impressions of tergites III–V coarse ( Fig. 91 ). Median lobe of aedeagus with conspicuously slender and acute apex in lateral view ( Figs 150–151 ). Distribution: Central Nepal ( Map 3 ). ................ truncatus

22. Anterior impressions of tergites III–V with few fine non-setiferous punctures confined to the middle. Species of rather large size with a rather large, but weakly transverse pronotum (approximately 1.1 times as broad as long). ............................................................................................................................................................................................ 23

– Anterior impressions of tergite III–V with more numerous non-setiferous punctures everywhere. .................. 24

23. Legs of uniformly pale-brown coloration. Pronotum sparsely and distinctly irregularly punctate; lateral margins not sinuate near posterior angles ( Fig. 43 ). Punctation of elytra coarse and irregularly distributed, anteriorly dense and somewhat asperate, posteriorly very sparse ( Fig. 43 ). Median lobe of aedeagus as in Figs 121–122 . Distribution: Northwest India ( Map 1 ). ..................................................................................................................... exasperatus

– Legs bicoloured, with dark-brown to blackish-brown femora and with paler tibiae and tarsi. Pronotum with nearly regularly distributed punctation ( Fig. 44 ). Elytra with much finer, much denser, and not distinctly irregularly distributed punctation, only slightly sparser posteriorly than anteriorly ( Fig. 44 ). Median lobe of aedeagus as in Figs 125–129 . Distribution: North India, Nepal ( Map 2 ). ............................................................................. pindarae

24. Small species; body length 4.3–5.0 mm; length of forebody 1.9–2.1 mm. Antennomere XI conspicuously elongate, approximately as long as the combined length of VIII–X ( Fig. 27 ). Head relatively large in relation to pronotum ( ASSING 2016a: figure 97). Aedeagus: ASSING (2016a: figures 272–273). Distribution: Oriental and southern East Palaearctic regions ( Map 5 ). ............................................................................................................................. castaneus

– Distinctly larger species. Antennomere XI less elongate, shorter than the combined length of VIII–X. ........... 25

25. Anterior third to anterior half of elytra or only humeral portions yellowish, posterior half (including suture) blackish ( Fig. 45 ). Aedeagus as in Figs 132–133 . Distribution: North India, Nepal ( Map 3 ). ............... perforatus

– Elytra yellowish with the postero-lateral portions more or less triangularly blackish; yellowish coloration extending posteriad along suture to posterior margin, or nearly so ( Figs 46–47 ). ................................................................... 26

26. Tergites VII–X reddish ( Fig. 87 ). Distribution: North India ( Map 1 ). ...................................................... championi

– Tergite VII blackish; posterior portion of tergite VIII somewhat infuscate ( Fig. 88 ). Aedeagus as in Figs 136–137 . Distribution: Nepal ( Map 1 ). ............................................................................................................................... pallipes

3.3 Key to the species of the subgenus Zyras recorded from Thailand

Up to today, 13 named species of Zyras sensu strictu have been recorded from Thailand, four of them exclusive. In addition, several unnamed species have been examined, one of them represented by a male without antennae and the remainder represented exclusively by females.

1. Pronotum red. ...................................................................................................................................................................... 2

– Pronotum dark-brown to black. ........................................................................................................................................ 5

2. Head red, of similar coloration as pronotum ( Fig. 61 ). Elytra with very coarse and dense punctation ( Fig. 61 ). Aedeagus as in Figs 187–188 . Distribution: Thailand, Malaysia. .................................................................... russiceps

– Head blackish. ...................................................................................................................................................................... 3

3. Very large and robust species; length of forebody> 3.5 mm; width of pronotum approximately 1.5 mm. Elytra reddish with the postero-lateral portion more or less distinctly and more or less extensively infuscate. Legs usually uniformly yellowish. Median lobe of aedeagus as in Figs 276–277 . Distribution: Oriental and southern East Palaearctic regions ( Map 10 ). ................................................................................................................................ preangeranus

– Smaller and more slender species; length of forebody <3.5 mm; width of pronotum <1.2 mm. Elytra blackish, often with slight bluish hue. At least the apices of the femora usually infuscate. ....................................................... 4

4. Profemora and the apical halves of the meso- and metafemora blackish-brown to black. Antennae very slender, approximately 2.7 mm long; antennomeres IV–VII distinctly oblong and X very weakly transverse. Elytra with rather coarse punctation. Distribution: Thailand ( Map 8 ). ............................................................................ thaiorum

– Femora yellowish with the apices usually narrowly infuscate. Antennae much shorter, <2.3 mm long; antennomeres V–VII not oblong and X distinctly transverse. Elytra with very fine punctation. Aedeagus: ASSING (2016a: figures 274–281). Distribution: Oriental and southern East Palaearctic regions ( Map 8 ). ........................... geminus

5. Legs bicoloured, at least the apices of the meso- and metafemora more or less distinctly infuscate. ...................... 6

– Legs uniformly pale-yellowish. .......................................................................................................................................... 7

6. Apices of meso- and metafemora narrowly and weakly infuscate. Elytra short and with fine punctation ( Fig. 70 ); hind wings reduced. Segments II–V of abdomen reddish ( Fig. 107 ). Aedeagus: Figs 212–215 . Distribution: South Thailand ( Map 10 ). .............................................................................................................................................. ambulans

– Profemora dark-brown; apical halves of meso- and metafemora blackish. Elytra longer and with coarser punctation ( Fig. 66 ). Most of segments II–V blackish ( Fig. 103 ). Male unknown. Distribution: Thailand, Southwest China ( Map 8 ). ................................................................................................................................................................... brignolii

7. Anterior abdominal tergites of dark coloration (posterior margins of segments may be paler). ............................. 8

– At least tergites II–V reddish. .......................................................................................................................................... 10

8. Antennae massive; antennomere XI barely as long as the combined length of IX and X ( ASSING 2016a: figure 41). Pronotum distinctly transverse, at least 1.2 times as broad as long and approximately 1.3 times as broad as head, rather finely and regularly punctate. Median lobe of aedeagus: ASSING (2016a: figures 215–216). Distribution: Thailand, China: Yunnan ( Map 8 ). ............................................................................................................... caloderoides

– Antennae slender; antennomere XI conspicuously elongate, approximately as long as the combined length of VIII–X ( Figs 28–29, 33 ). Pronotum slender, much less than 1.2 times as broad as long and only slightly broader than head ( Figs 65 , 69 ); punctation coarse and irregularly distributed. ........................................................................................ 9

9. Antennae more slender ( Figs 28–29 ). Pronotum with very uneven surface (i.e., with more or less irregular impressions); punctation conspicuously irregularly distributed ( Fig. 65 ). Male tergite VIII with four blunt projections posteriorly ( Fig. 196 ). Median lobe of aedeagus as in Figs 191–194 . Distribution: Oriental and southern East Palaearctic regions ( Map 6 ). ................................................................................................................................ proximus

– Antennae less slender ( Fig. 33 ). Pronotum without impressions and with less irregularly distributed punctation ( Fig. 69 ). Male tergite VIII only with a median pair of less distinct projections posteriorly ( Fig. 206 ). Median lobe of aedeagus as in Figs 208–211 . Distribution: Thailand ( Map 10 ). .............................................................. parvicollis

10. Antennomere XI conspicuously elongate, approximately as long as the combined length of VIII–X ( Figs 27, 30–31 ). .............................................................................................................................................................................................. 11

– Antennomere XI less elongate, shorter than the combined length of VIII–X. ......................................................... 12

11. Pronotum with uneven surface, more or less irregularly impressed; punctation coarse and irregularly distributed ( Fig. 67 ). Anterior impressions of tergites III–V and anterior portion of tergite VI with coarse and dense nonsetiferous punctation ( Fig. 106 ). Posterior margin of male tergite VIII with four blunt projections ( Fig. 201 ). Median lobe of aedeagus as in Figs 198–199 . Distribution: Thailand, Laos ( Map 5 ). ........................................... novinversus

– Pronotum with smooth surface, without impressions; punctation fine and more or less regularly distributed ( ASSING 2016a: figure 97). Anterior impressions of tergites III–V and anterior portion of tergite VI with sparse and very fine non-setiferous punctation ( ASSING 2016a: figure 154). Posterior margin of male tergite VIII with shallow concavity in the middle. Median lobe of aedeagus: ASSING (2016a: figures 272–273). Distribution: Oriental and southern East Palaearctic regions ( Map 5 ). ....................................................................................................................... castaneus

12. Pronotum strongly transverse, nearly 1.3 times as broad as long and 1.2 times as broad as head ( Fig. 68 ). Tergite VII nearly completely blackish-brown to black ( Fig. 102 ). Median lobe of aedeagus with very short ventral process ( Figs 203–204 ). Distribution: Thailand ( Map 10 ). ..................................................................................... brevilobatus

– Pronotum weakly transverse, approximately 1.15 times as broad as long ( ASSING 2016a: figure 172). Tergite VII bicoloured with the anterior third reddish and the posterior two-thirds black ( ASSING 2016a: figure 174). Median lobe of aedeagus: ASSING (2016a: figures 231–232). Distribution: Oriental and southern East Palaearctic regions ( Map 9 ). ................................................................................................................................................................ bettotanus

Figures

Map 1: Revised distributions of Zyras kraatzi (black and white circles; white circle: type locality), Z. exasperatus (white circle), Z. iniquus (triangles), Z. pallipes (diamonds), and Z. championi (white stars) in the Himalaya.

Map 2: Revised distributions of Zyras pindarae (black circles) and Z. nigroaeneus (white circles) in the Himalaya.

Map 3: Revised distributionsof Zyras perforatus (blackcircles),Z. morvani(white circles),andZ. truncatus(white star)in theHimalaya.

Map 4: Revised distributions of Zyras condignus (black circles) and Z. morulus (white circles) in the Himalaya.

Map 5: Revised distributions of Zyras castaneus (black circles) and Z. novinversus (white circles) in the southern East Palaearctic

Map 6: Revised distributions of Zyras geminus (black circles) and Z. proximus (white circles) in the southern East Palaearctic and Oriental regions.

Map 7: Revised distributions of Zyras gardneri (black circles), Z. hirsutiventris (white diamonds), Z. parageminus (black triangles), Z. hastatus (white triangle), Z. hirtus (white circles), and Z. latilobatus (star) in India and adjacent regions.

Map 8: Revised distributions of Zyras alternans (black circles), Z. luteipes (star), Z. brignolii (white circles), and Z. nitens (white

Map 9: Revised distributions of Zyras bettotanus (black circles), Z. nilgiriensis (star), and Z. longilobatus (white circles) in the

Map 10: Revised distributions of Zyras preangeranus (black circles), Z. brevilobatus (white star), Z. parvicollis (white star), Z. ambulans (white circle), Z. densissimus (black star), Z. titan (black star), Z. densihirtus (black star), and Z. nigrihirtus (black star) in the southern East Palaearctic and Oriental regions.

Figs 55–67: Forebody of Zyrasspp.:parageminus (55–56; 55:holotype;56:holotype ofZ. nameriensis),latilobatus (57),alternans (58), hirtus (59), nilgiriensis (60), russiceps (61), gardneri (62), hirsutiventris (63),luteipes (64), proximus, holotype (65), brignolii, holotype of Z. thainiger (66), novinversus (67). Scale bar: 1.0 mm.

Figs 82–99: Abdomen of Zyras spp.: pindarae (82–85), perforatus (86), championi (87), pallipes (88), longilobatus (89), hastatus (90), truncatus (91), latilobatus (92), morvani (93), condignus (94), hirtus (95), gardneri (96), hirsutiventris (97), parageminus (98), alternans (99). Scale bar: 1.0 mm.

Figs 1–41:Antennae of Zyrasspp.:kraatzi(1),exasperatus(2),pindarae (3–5),perforatus(6),championi(7),pallipes (8),condignus (9), nitens (10), truncatus (11), nilgiriensis (12), hirsutiventris (13), russiceps (14), brignolii (15), hastatus (16), parageminus (17–18), latilobatus (19), alternans (20), hirtus (21), nigroaeneus (22), luteipes (23), longilobatus (24), morvani (25), gratellus (Malaysia) (26), castaneus, lectotype (27),proximus (28–29),novinversus (30–31), brevilobatus(32), parvicollis(33), ambulans (34),matangensis (35), densissimus (36), densihirtus (37), nigrihirtus (38), gratellus (Sulawesi) (39), gardneri (40), titan (41). Scale bar: 1.0 mm.

Figs 174–190: Zyras gardneri (174–177), Z. hirsutiventris (178–181), Z. luteipes (182–186), and Z. russiceps (187–190): median lobe of aedeagus in lateral and in ventral view (174–175, 178–179, 182–183, 187–188); male tergite VIII (176, 180, 185, 189); male sternite VIII (177, 181, 186, 190); paramere (184). Scale bars: 0.5 mm.

Figs 100–116: Abdomen of Zyras spp.: luteipes (100), nilgiriensis (101), brevilobatus(102), brignolii (103), parvicollis (104), proximus, holotype (105), novinversus (106), ambulans (107), matangensis (108), nitens (109), gratellus (Malaysia) (110), densissimus (111), gratellus (Sulawesi) (112), titan (113), russiceps (114), densihirtus (115), nigrihirtus (116). Scale bar: 1.0 mm.

Figs 156–173: Zyras longilobatus (156–160), Z. latilobatus (161–165), Z. alternans (166–169), and Z. hirtus (170–173; 170–171: Sri Lanka; 172–173: South India): median lobe of aedeagus in lateral and in ventral view (156–157, 161–162, 166–167, 170–173); paramere (158, 163); male tergite VIII (159, 164, 168); male sternite VIII (160, 165, 169). Scale bars: 0.5 mm.

Figs 42–54: Forebody of Zyras spp.: kraatzi (42), exasperatus (43), pindarae (44), perforatus (45), championi (46), pallipes (47), condignus, holotype (48), condignus, Nepal (49), nigroaeneus (50), morvani (51), truncatus (52), longilobatus (53), hastatus (54). Scale bar: 1.0 mm.

Figs 260–277: Zyras morulus (260–265), Z. variolatus (266–270), Z. montanus (271–275), and Z. preangeranus from Java (276–277): forebody (260); antenna (261); abdomen (262); median lobe of aedeagus in lateral and in ventral view (263–264, 266–267, 271–272, 276–277); paramere (265, 268, 273); male tergite VIII (269, 274); male sternite VIII (270, 275). Scale bars: 260–262: 1.0 mm; 263–277: 0.5 mm.

Figs 117–133: Zyras kraatzi (117–120), Z. exasperatus (121–124), Z. pindarae (125–131), and Z. perforatus (132–133): median lobe of aedeagus in lateral and in ventral view (117–118, 121–122, 125–128, 132–133); male tergite VIII (119, 123, 130); male sternite VIII (120, 124, 131); paramere (129). Scale bars: 0.5 mm.

Figs 68–81: Forebody (68–77), head and pronotum (78), and abdomen (79–81) of Zyras spp.: brevilobatus (68), parvicollis (69), ambulans (70), nitens, holotype (71), gratellus (Malaysia) (72), matangensis (73), gratellus (Sulawesi) (74), densissimus (75), titan (76), densihirtus (77), nigrihirtus (78), kraatzi (79), asperatus (80), nigroaeneus (81). Scale bar: 1.0 mm.

Figs 134–155: Zyras perforatus (134–135), Z. pallipes (136–139), Z. nigroaeneus, holotype (140–141), Z. condignus (142–147), Z. morvani, holotype (148–149), and Z. truncatus (150–155): male tergite VIII (134, 138, 140, 146, 148, 152); male sternite VIII (135, 139, 141, 147, 149, 153); median lobe of aedeagus in lateral and in ventral view (136–137, 142–143, 150–151); paramere (144–145); female tergite VIII (154); female sternite VIII (155). Scale bars: 0.5 mm.

Figs 191–207: Zyras proximus (191–197; 191–192: Thailand; 193–197: India), Z. novinversus (198–202), Z. brevilobatus (203–205), and Z. parvicollis (206–207): median lobe of aedeagus in lateral and in ventral view (191–194, 198–199, 203–204); paramere (195, 200, 205); male tergite VIII (196, 201, 206); male sternite VIII (197, 202, 207). Scale bars: 0.5 mm.

Figs 208–226: Zyras parvicollis (208–211), Z. ambulans (212–215), Z. nitens, holotype (216–221), and Z. gratellus (Malaysia) (222–226):medianlobeofaedeagusinlateralandinventralview(208–213,216–217,222–223);paramere (214,218,224); male tergite VIII (219, 225); male sternite VIII (220, 226); apical lobe of paramere (215); postero–median portion of male sternite VIII (221). Scale bars: 208–214, 216–220, 222–226: 0.5 mm; 215, 221: 0.1 mm.