Tylopus, Jeekel, 1968

Likhitrakarn, Natdanai, Golovatch, Sergei I. & Panha, Somsak, 2016, The millipede genus Tylopus Jeekel, 1968 (Diplopoda, Polydesmida, Paradoxosomatidae), with a key and descriptions of eight new species from Indochina, European Journal of Taxonomy 195, pp. 1-47 : 39-45

Key to the known species of Tylopus (based chiefly on Ƌ), modified after Likhitrakarn et al. (2010):

1. Most of Ƌ prefemora evidently swollen laterally ( Figs 13C , 24C ) ……………………………2 – All Ƌ prefemora normal, not bulged laterally ( Figs 2G , 11F, G , 17 E–G, 19C) …………29

2. Surface of metaterga virtually smooth, at best extremely faintly rugulose in certain places and/ or with a few barely traceable (setigerous) tubercles near caudal margin (setae mostly broken off) ( Figs 7B, D , 12A – B, D , 21A, C ) ……………………………………………………………………3

– Surface of metaterga mostly rugulose to coarsely rugose/tuberculate ( Figs 18A – B, D , 23B, D ) ………………………………………………………………………………………………………9

3. Paraterga moderately developed ( Figs 1A, C , 21A, C ), ratio of Ƌ midbody prozonite to metazonite width below 1:1.2 …………………………………………………………………………………4

– Paraterga relatively well-developed ( Figs 10B, D , 11B, D , 18B, D ), radio of Ƌ midbody prozonite to metazonite width over 1:1.2 …………………………………………………………5

4. Transverse sulcus on metaterga starting with segment 5. Calluses without incisions. Gonopod with process h smallanddentiform. Thailand ……………………… T.prosperus Golovatch&Enghoff,1993

– Transverse sulcus on metaterga starting with segment 4. Calluses with faint one or two denticles on poreless and poriferous paraterga, respectively. Gonopod very simple, without processes. Southern China ………………………………………………………… T. reductus Golovatch, 2013

5. Calluses without incisions. Gonopod postfemoral lobe l much broader than long; area basal to l delimited by a distinct cingulum ……………………………………………………………………6

– Calluses mostly with 1–2 incisions. Gonopod postfemoral lobe l either as long as broad or longer; no cingulum basal to l ………………………………………………………………………………7

6. Body smaller: length up to ca 19–30 mm. Gonopod with a broadly coiled solenophore and a small finger-shaped process on lobe l. China …………………………… T. kabaki Golovatch, 2014

– Body larger: length ca 33–35 mm. Gonopod with a slender and coiled solenophore, but without processes on lobe l. Vietnam …………………………………………… T. magicus Golovatch, 1984

7. Metaterga at best only very faintly rugulose near stricture, near sulcus and/or at base of paraterga, with 2–3 weak, oblong tubercles near rear margin. Ƌ postfemora and tibiae tuberculiferous. Gonopods with process z serrate, with 3 spines. Thailand ………… T. similirugosus Golovatch & Enghoff, 1993

– Metatergal surface entirely smooth, polished, without tubercles. Ƌ legs without adenostyles (= tubercles). Gonopods with process z pointed, without spines ……………………………8

8. Midline wanting. Gonopods with process h without additional tooth, while process z very high. Vietnam ………………………………………………………………… T. mutilatus ( Attems, 1953)

– Midline mostly traceable on prozonae and anterior halves of metaterga. Gonopods with process h higher with an evident dorsal tooth, while process z small( Figs13A – B , 14 – 15 ). Laos ………………… ……………………………………………………………………………… T. acuminatus sp. nov.

9. Body larger: 37–42 mm long, 2.8–3.8 and 4.1–5.0 mm wide on pro- and metaterga, respectively …… …………………………………………………………………………………………………10 – Body smaller ………………………………………………………………………………………15

10. Sterna between Ƌ coxae 4 with a pair of separated cones. Gonopod with a short spiniform process h, a basally only poorly delimited lobe l and a small lobe-shaped process z. Thailand ………………………………… T. grandis Likhitrakarn, Golovatch, Prateepasen & Panha, 2010

– Sterna between Ƌ coxae 4 with a slightly notched sternal lobe. Gonopod with a rather long process h carrying two branches, plus a basally evidently delimited lobe l and a large, serrated process z. Vietnam ……………………………………………………………… T. phanluongi Nguyen, 2012

11. Both processes h and z of gonopod spiniform. Thailand …………………………………………… ………………………………… T. bispinosus Likhitrakarn, Golovatch, Prateepasen & Panha, 2010 – Gonopod otherwise ………………………………………………………………………………12

12. Gonopod process h subflagelliform, process m extremely long and prominent. Thailand ………… ………………………………… T. extremus Likhitrakarn, Golovatch, Prateepasen & Panha, 2010 – Gonopod otherwise ………………………………………………………………………………13

13. Gonopod process h prominent, spine s high, rather slender, curved, nearly pointed. Vietnam …………………………………………………………………………… T. sapaensis Nguyen, 2012 – Gonopod otherwise ………………………………………………………………………………14

14. Ƌ legs shorter, ca 1.2–1.3 times as long as midbody height. Gonopod lobe l velum-shaped and supplied with two denticles; spine z short and knife-shaped, while spine h rudimentary. Thailand ………………………………… T. veliger Likhitrakarn, Golovatch, Prateepasen & Panha, 2010

– Ƌ legs longer, ca 1.6–1.7times as long as midbody height.Gonopod spine z small, placed closer to base of spine h. Thailand ………… T. parajeekeli Likhitrakarn, Golovatch, Prateepasen & Panha, 2010

15. Most of metaterga with a pattern of 2+2 and 2+2 setigerous tubercles in two rows, rear row somewhat less strongly developed than fore one. Myanmar and Thailand … T. doriae ( Pocock, 1895)

– Most of metaterga with rear row of setigerous tubercles or wrinkles more strongly developed than fore row, the latter (next to) wanting ………………………………………………………………16

16. Transverse sulcus on metaterga starting with segment 4, either fully or almost fully developed there, always fully developed starting with segment 5 ……………………………………………………17 – Transverse sulcus on metaterga starting only with segment 5 ……………………………………20

17. Transverse sulcus fully developed and reaching base of paraterga already starting with segment 4. Gonopod tooth z at base of lobe l coarsely serrate along proximal margin. Vietnam ………………… ……………………………………………………………………………… T. hilaris ( Attems, 1937)

– Transverse sulcus fully developed only starting with segment 5. Gonopod tooth z either devoid of serration or serrate along distal margin ……………………………………………………………18

18. Paraterga 2 rather broadly rounded caudolaterally. Gonopod relatively simple, process h poorly developed, no additional outgrowths near base. Thailand ………………………………………… ……………………………………………………………… T. affinis Golovatch & Enghoff, 1993

– Paraterga 2 pointed caudally. Gonopods more complex …………………………………………19

19. Coloration dark brown, without a cingulate pattern. Sternal lamina between Ƌ coxae 4 low and distinctly bimodal. Gonopod tooth z prominent and serrate along distal margin. Thailand …………………………………………………………… T. rugosus Golovatch & Enghoff, 1993

– Coloration pale, with a cingulate pattern. Sternal lamina between Ƌ coxae 4 high, subquadrate. Gonopod tooth z smaller and spiniform. Thailand ……… T. semirugosus Golovatch & Enghoff, 1993

20. Paratergal corner protruding caudad behind rear margin only starting with segment 15, being obtuse-angled or subrectangular and lying more or less within the margin until segment 14. Vietnam ……………………………………………………………… T. hilaroides Golovatch, 1984

– Paratergal corner protruding caudad behind rear margin before segment 14, mostly pointed …21

21. Pattern of tergal setation on segments 18 and/or 19: 2+2 and 5+ 5 in two rows ……………22 – Pattern of tergal setation at least on segments 5–19: 2+2 and 4+ 4 in two rows ……………25

22. Pattern of tergal setation 2+2 and 5+5 on both segments 18 and 19. Paraterga 2 pointed caudally. Epiproct with pre-apical incisions very close to apical knobs. Sternal lamina between Ƌ coxae 4 an unusually low and even ridge. Adenostyles on midbody Ƌ postfemora and, to a lesser extent, tibiae exceptionally prominent. Thailand …… T. poolpermorum Golovatch & Enghoff, 1993

– Pattern of tergal setation 2+2 and 5+5 on segment 19. Paraterga 2 more or less narrowly rounded. Pre-apical incisions on epiproct more strongly removed from tip. Sternal lamina between Ƌ coxae 4 concave medially. Ventral adenostyles on Ƌ legs less prominent ……………………23

23. Body smaller: width ca 2.0 mm. Sternal lamina between Ƌ coxae 4 a pair of separated setigerous tubercles. Ventral adenostyles on Ƌ legs almost missing. Gonopods without any outgrowths near base of process h. Thailand ………………………………… T. haplorugosus Golovatch & Enghoff, 1993

– Body larger: width over 3.0 mm. Sternal lamina between Ƌ coxae 4 single. Ventral adenostyles on Ƌ legs more prominent. Gonopod with a spine near base of process h ………………………24

24. Sternal lamina between Ƌ coxae 4 high, emarginated. Adenostyles on Ƌ postfemora and tibiae well-developed. Gonopods rather simple, spine z inconspicuous. Thailand ……………………… …………………………………………………………… T. allorugosus Golovatch &Enghoff, 1993

– Sternal lamina between Ƌ coxae lower, slightly concave. Adenostyles on Ƌ postfemora and tibiae less strongly developed. Gonopods more complex, spine z long and large. Thailand ……………………………………………………………………… T. perarmatus Hoffman, 1973

25. Paraterga 2 pointed caudally. Sternal lamina between Ƌ coxae 4 exceptionally densely setose, low and concave ventrally. Gonopods with a medium-sized process h and a smaller lobular z at base of h. Thailand ……………………………………………………. T. jeekeli Golovatch & Enghoff, 1993

– Paraterga 2 more or less narrowly rounded caudally. Sternal lamina between Ƌ coxae 4 higher and less strongly setose. Gonopod outgrowths h and z either almost wanting or very large …26

26. Sternal lamina between Ƌ coxae 4 with a straight ventral margin. Pleurosternal carinae poorly developed, in Ƌ slightly drawn behind rear margin only until segments 8–10 …………………27

– Sternal lamina between Ƌ coxae 4 slightly concave ventrally. Pleurosternal carinae better developed, in Ƌ slightly drawn behind rear margin at least until segment 15 …………………28

27. Body smaller: width up to 3.1–3.2 mm. Mid-dorsal line very clear on both halves of metaterga. Gonopods relatively simple, with both h and z almost wanting. Thailand ………………………… …………………………………………………………… T. hoffmani Golovatch & Enghoff, 1993

– Body larger: width 4.0– 5.3 mm. Mid-dorsal line not so well-developed at least on rear halves of metaterga. Gonopods more complex, with both h and z very conspicuous ( Figs 24A – B , 25 ). Thailand ……………………………………………… T. baenzigeri Golovatch & Enghoff, 1993

28. Metatergum 19 slightly rugulose posteriorly. Calluses on segment 2 with three, on following paraterga with two, incisions. Gonopods extremely complex, with numerous spiniform outgrowths. Thailand ………………………………………………………… T. perplexus Golovatch & Enghoff, 1993

– Metatergum 19 entirely smooth. Calluses with two or three incisions on poreless and poriferous paraterga, respectively. Gonopod less strongly differentiated. Thailand ……………………………… ……………………………………………………………… T. amicus Golovatch & Enghoff, 1993

29. Either most of Ƌ sterna with an oblique tubercle or spine, or only anterior sterna with a small cone, near coxae …………………………………………………………………………………………30 – Neither spines nor tubercles on Ƌ sterna …………………………………………………………37

30. Calluses virtually devoid of incisions………………………………………………………………31 – Calluses always at least with one lateral incision …………………………………………………32

31. Ƌ coxae 4 with a pair of separated cones. Gonopod process h small, while solenophore slender and short. China …………………………………………………………… T. sinensis Golovatch, 1995

– Ƌ coxae 4 with an entire, subtrapeziform, sternal lobe. Gonopod process h larger, while solenophore stout and long. Vietnam ………………………………………………… T. golovatchi Nguyen, 2012

32. A pair of small, fully separated, setose cones between Ƌ coxae 4 ( Fig. 10I – J ). Gonopod with process h rather short, slightly flattened, blunt ( Fig. 11 A–D). Laos ………… T. retusus sp. nov. – An entire lobe between Ƌ coxae 4. Gonopod otherwise …………………………………………33

33. Metaterga strongly rugose/tuberculate/granulate ( Fig. 18 A–F, H) ………………………………34 – Metaterga entirely smooth and polished, devoid of evident tubercles, at best extremely faintly rugulose near transverse sulcus ( Fig. 1 A–F) ……………………………………………………35

34. Calluses mostly with one lateral, clear incision. Pleurosternal carinae missing after segment 17. Gonopod processes h, z, m and spine x present ( Fig. 19A – B ). Laos … T. thunghaihin sp. nov.

– Calluses with two lateral setigerous incisions. Pleurosternal carinae missing after segment 12. Only gonopod process h present. Vietnam …………………………… T. spinisterna Nguyen, 2012

35. Only rear sternum on most of Ƌ segments with a pair of small spines. Metaterga mostly with 2+2 and 3+3 setigerous tubercles in two transverse rows. Pleurosternal carinae missing after segment 9. Myanmar …………………………………………………………… T. silvestris ( Pocock, 1895)

– Fore and rear sterna of most of Ƌ segments with a pair of small tubercles and spines, respectively. Pleurosternal carinae present until segment 18 ……………………………………………………36

36. Metaterga with four transverse rows of very small setigerous tubercles or setae. Ƌ tarsal brushes present until leg 6. Ƌ femora 5 with a very strong, distoventral, densely pilose adenostyle ( Fig. 2F ). Gonopod process h higher and pointed ( Fig. 2 A–D). ( Fig. 26 ). Thailand …… T. flavolineatus sp. nov.

– Metaterga with two transverse rows of small tubercles or spines, respevtively Ƌ tarsal brushes missing. Ƌ legs normal. Gonopod process h very small. Vietnam …………… T. maculatus Golovatch, 1984

37. Paraterga virtually missing ( Figs 4 A–H, 7A–H) …………………………………………………38 – Paraterga well- or strongly developed ( Figs 1A, C , 10B, D , 12B, D , 16B, D , 18B, D , 23B, D ) …39

38. Antennae shorter ( Fig. 4A ), drawn behind only body segment 2 when stretched dorsally. A small, rounded, densely setose, sternal cone between Ƌ coxae 4 ( Fig. 4I – J ). Goponod process h prominent, while process z long and rather slender ( Figs 5 – 6 ). Thailand ……… T. hongkhraiensis sp. nov.

– Antennae longer ( Fig. 7A ), reaching behind body segment 3. An entire, high, inverted funnel-shaped, sternal lobe between each of Ƌ coxae 3 and 4 ( Fig. 7I – J ). Goponod without processes ( Figs 8 – 9 ). Laos ……………………………………………………………………………… T. moniliformis sp. nov.

39. Metaterga entirely smooth and polished, devoid of evident tubercles, at best extremely faintly rugulose near transverse sulcus ……………………………………………………………………40

– Metaterga rather clearly rugose/tuberculate/granulate, posterior row of setae at least partly borne on tubercles……………………………………………………………………………………………46

40. Transverse sulcus on metaterga starting with segment 3 or 4, but fully developed and reaching bases of paraterga starting with segment 4 or 5. Gonopod process h large ( Fig. 17 A–D) ………41 – Transverse sulcus on metaterga starting only with segment 5. Gonopod process h small ( Fig. 22 ) …44

41. Gonopod with a distinct sharp process on lobe l. Ventral adenostyles on Ƌ legs absent. Southern China ………………………………………………………………………… T. similis Golovatch, 2014

– Gonopod without processes on lobe l. Ventral adenostyles on Ƌ legs: a distal knob on femur, a distomedial knob on postfemur, and a parabasal knob on both tibia and tarsus ……………42

42. Head wider than collum and subequal in width to segment 3. Gonopod process h at about midlength with a strong ventral outgrowth, while solenophore without processes. Vietnam ……… ……………………………………………………………………… T. procurvus Golovatch, 1984

– Head narrower than collum. Gonopod process h without outgrowths, while process x present on solenophore ………………………………………………………………………………………43

43. Antennae shorter ( Fig. 16A ), extending only behind body segment 2 when stretched dorsally. Ƌ tarsal brushes present until segment 6. Gonopod process x longer, the latter’s tip rounded dorsally ( Fig. 17 A–D). Laos …………………………………………………… T. dorsalis sp. nov.

– Antennae longer, reaching until body segment 5. Ƌ tarsal brushes present on all segments. Gonopod process x shorter, its tip pointed. Vietnam …………………………… T. crassipes Golovatch, 1984

44. Small species: width 1.6 mm. Only a single row of 2+2 tergal setae. Metaterga entirely smooth. Adenostyles on Ƌ legs present. Vietnam …………………………………… T. sigma ( Attems, 1953)

– Body wider than 2.0 mm. Metaterga with two transverse rows of setae or traceable insertion points. Metaterga very finely rugulose only near transverse sulcus. Adenostyles on Ƌ legs absent or represented by pads in their stead ……………………………………………………………45

45. Paraterga broader: 3.1 mm wide. Pleurosternal carinae present until segment 13 or 14. Pads present on Ƌ femora, postfemora, tibiae (all distally) and tarsi (almost entirely). Thailand ……………… …………………………………………………………… T. pulvinipes Golovatch & Enghoff, 1993

– Paraterga narrower: 2.1–2.5 mm wide. Pleurosternal carinae present until segment 17. Ƌ legs without pads. Myanmar ………………………………………………………… T. punctus sp. nov.

46. Metatergal surface polished and smooth except for two rows of conspicuous tubercles ………47 – Metaterga at least partly rugulose/rugose to granular; at most one row of tubercles ……………52

47. Ƌ legs without adenostyles (= tubercles). Gonopods with process h higher than solenophore, strongly twisted, pointed apically. Southern China ………………… T. schawalleri Golovatch, 2013 – Ƌ legs with adenostyles (= tubercles) at least on segment 6. Gonopods otherwise. Vietnam …48

48. Paraterga very poorly developed, rounded, low, projecting slightly behind rear margin like small knobs only on segments 18 and 19. Calluses virtually devoid of incisions. Transverse sulcus on metaterga poorly developed, starting already with segment 2, although fully developed only starting with segment 5. A paramedian pair of denticles between Ƌ coxae 5 behind a prominent, subquadrate lamina between Ƌ coxae 4. Gonopod process h entirely missing. Thailand ……………………………………………… T. strongylosomoides (Korsós & Golovatch, 1989)

– Paraterga better developed, protruding behind rear margin at least starting with segment 5. Calluses always at least with one lateral incision. Transverse sulcus starting only with segments 3–5. Dentiform tubercles between Ƌ coxae 5 missing. Gonopod process h invariably present ………49

49. Body larger:more than 25 mm long.Ƌ tarsal brushes not present on all legs.Gonopod lobe l normal…50 – Body smaller: less than 18 mm long. Ƌ tarsal brushes present on all legs. Gonopod lobe l with a spine apically ………………………………………………………………………………………51

50. Ƌ tarsal brushes present on legs 1–15. Pleurosternal carinae present until segment 9. Gonopod process h larger, finger-shaped. Vietnam ………………………… T. roseiparaterga Nguyen, 2012

– Ƌ tarsal brushes present on legs 5–30. Pleurosternal carinae present until segment 17. Gonopod process h small, slender and pointed. Southern China ……………… T. deharvengi Liu & Luo, 2013

51. Paraterga acutangular caudally and pointed beak-like already starting with collum. Tergal setigerous tubercles: 3+3 and 5+5 on segments 16–19. Adenostyle pattern on Ƌ legs: a distal knob on femora and a parabasal knob on most of postfemora, tibiae and tarsi. Gonopod process h large, lamellar and sigmoid. Vietnam …………………………………………………… T. granulatus Golovatch, 1984

– Paraterga acutangular caudally and pointed beak-like only starting with segment 4. 2+2 and 4+4 tergal setigerous tubercles on segments 16–19. Adenostyle pattern on Ƌ legs: a proximal fingershaped tubercle crowned with a bunch of setae only on femora 6, 8 and 9. Gonopod process h smaller, spiniform. Vietnam ……………………………………………………… T. topali Golovatch, 1984

52. Transverse sulcus on metaterga starting and fully developed only with segment 5. Ventral adenostyles present on all Ƌ podomeres except coxa. Vietnam and/or Laos ……………53

– Transverse sulcus on metaterga starting with segment 4, but fully developed only starting with segment 5. Ventral tubercles only on some of Ƌ telopoditomeres. Thailand ………………54

53. Metaterga rugulose also in front of transverse sulcus, at rear margin with several oblong tubercles. Sternal lamina between Ƌ coxae 4 a pair of setigerous knobs. Neither gonopod lobe m nor lobe l spinigerous. Vietnam ………………………………………… T. tamdaoensis Korsós & Golovatch, 1989

– Metaterga rugose only behind transverse sulcus, without evident tubercles at rear margin. Sternal lamina between Ƌ coxae 4 single. Both gonopod lobe m and lobe l crowned with a spine. Laos and Vietnam …………………………………………………………… T. nodulipes ( Attems, 1953)

54. Metaterga modestly rugulose only near transverse sulcus, posteriorly neither granular nor microtuberculate. Calluses broad. Gonopod process h simple, high, never particularly coiled; lobe l very moderately serrate at apex ……………………………………………………………………55

– Metaterga distinctly rugose-granular/microtuberculate even on fore halves. Calluses narrow. Gonopod process h better developed and more strongly coiled; lobe l apically either bare or with a digitiform outgrowth ……………………………………………………………………………56

55. Caudal corner of paraterga pointed starting with segment 3. Pleurosternal carinae particularly well-developed, drawn behind rear margin until segment 16 or 17. Adenostyles often present on Ƌ prefemora ………………………………………… T. degerboelae Golovatch & Enghoff, 1993

– Caudal corners of paraterga mainly narrowly rounded, pointed only starting with segment 15. Pleurosternal carinae less strongly developed ……………………… T. pallidus Golovatch & Enghoff, 1993

56. Coloration dark, brown. Sternal lamina between Ƌ coxae 4 a pair of separated setigerous tubercles preceded by another pair of very small tubercles between coxae 3. Gonopod lobe l devoid of an apical process ………………………………………………… T. asper Golovatch & Enghoff, 1993

– Coloration uniformly pale. Sternal lamina between Ƌ coxae 4 single, not accompanied by additional tubercles in front or behind. Gonopod lobe l with a strong, apical, finger-shaped process …… 57

57. Mostly 3+3 tubercles at rear margin of metaterga. Sternal lamina between Ƌ coxae 4 distinctly emarginate. Larger adenostyles close to midlength on Ƌ postfemora and tibiae, femora with a distal knob. Gonopod process h slenderer and shorter, apex of lobule m not developed into a spine ……………………………………………………… T. subcoriaceus Golovatch & Enghoff, 1993

– Mostly 4+4 tubercles at rear margin of metaterga. Sternal lamina between Ƌ coxae trapeziform. Larger adenostyles on both postfemora and tibiae more distal, femoral knob missing. Gonopod process h unusually prominent, with a hook at base; apex of lobule m spiniform ………………… …………………………………………………………… T. coriaceus Golovatch & Enghoff, 1993

Figures

Fig. 13. Tylopus acuminatus sp. nov., holotype, Ƌ, right gonopod. A–B. Lateral and mesal views, respectively. C. Leg 13. Scale bar: A–B. 0.5 mm; C = no scale bar. Abbreviations are explained in the text.

Fig. 24. Tylopus baenzigeri Golovatch & Enghoff, 1993, Ƌ. A–B. Right gonopod, lateral and mesal views, respectively. C. Leg 8. D. Legs of segment 11. Scale bars = 0.5 mm. Abbreviations are explained in the text.

Fig. 2. Tylopus flavolineatus sp. nov., holotype, Ƌ, left gonopod. A–D. Mesal, lateral, caudal and suboral views, respectively. E. Sterna of segment 10. F. Leg 6. G. Leg 10. Scale bars = 0.5 mm. Abbreviations are explained in the text.

Fig. 11. Tylopus retusus sp. nov., holotype, Ƌ, left gonopod. A–D. Mesal, lateral, subcaudal and suboral views, respectively. E. Sterna of segment 10. F. Leg 6. G. Leg 10. Scale bars = 0.2 mm. Abbreviations are explained in the text.

Fig. 17. Tylopus dorsalis sp. nov., holotype, Ƌ. A–D. Right gonopod, lateral, mesal, caudal and oral views, respectively. Scale bars = 0.2 mm. Abbreviations are explained in the text.

Fig. 7. Tylopus moniliformis sp. nov., holotype, Ƌ (B–J), paratypes, Ƌ, ♀ (A). A. Habitus, live coloration. B–C. Anterior part of body, dorsal and lateral views, respectively. D–E. Segments 10 and 11, dorsal and lateral views, respectively. F–H. Posterior part of body, subdorsal, ventral and sublateral views, respectively. I–J. Sternal cones between coxae 4, subcaudal and sublateral views, respectively. Abbreviations are explained in the text.

Fig. 12. Tylopus acuminatus sp. nov., holotype, Ƌ. A. Habitus, live coloration. B–C. Anterior part of body, dorsal and lateral views, respectively. D–E. Segments 10 and 11, dorsal and lateral views, respectively. F–H. Posterior part of body, dorsal, ventral and lateral views, respectively. I–J. Sternal cones between coxae 4, caudal and sublateral views, respectively.

Fig. 21. Tylopus punctus sp. nov., holotype, Ƌ. A–B. Anterior part of body, dorsal and lateral views, respectively. C–D. Segments 11–12, dorsal and lateral views, respectively. E–G. Posterior part of body, lateral, subdorsal and subventral views, respectively. H–I. Sternal cones between coxae 4, subcaudal and sublateral views, respectively.

Fig. 18. Tylopus thunghaihin sp. nov., holotype, Ƌ. A. Habitus, live coloration B–C. Anterior part of body, dorsal and lateral views, respectively. D–E. Segments 10 and 11, dorsal and lateral views, respectively. F–H. Posterior part of body, dorsal, ventral and lateral views, respectively. I–J. Sternal cones between coxae 4, caudal and sublateral views, respectively.

Fig. 23. Tylopus baenzigeri Golovatch & Enghoff, 1993, Ƌ. A. Habitus, live coloration. B–C. Anterior part of body, dorsal and lateral views, respectively. D–E. Segments 10 and 11, dorsal and lateral views, respectively. F–H. Posterior part of body, dorsal, ventral and lateral views, respectively. I–J. Sternal cones between coxae 4, subcaudal and sublateral views, respectively.

Fig. 1. Tylopus flavolineatus sp. nov., holotype, Ƌ (A, C–I), paratype, Ƌ (B). A–B. Anterior part of body, dorsal and lateral views, respectively. C–D. Segments 10 and 11, dorsal and lateral views, respectively. E–G. Posterior part of body, lateral, dorsal and subventral views, respectively. H–I. Sternal cones between coxae 4, caudal and sublateral views, respectively.

Fig. 10. Tylopus retusus sp. nov., holotype, Ƌ. A. Habitus, live coloration. B–C. Anterior part of body, dorsal and lateral views, respectively. D–E. Segments 10 and 11, dorsal and lateral views, respectively. F–H. Posterior part of body, dorsal, ventral and lateral views, respectively. I–J. Sternal cones between coxae 4, caudal and sublateral views, respectively.

Fig. 14. Tylopus acuminatus sp. nov., holotype, Ƌ, right gonopod. A–B. Mesal and lateral views, respectively. C–F. Distal part, sublateral, submesal, subcaudal and suboral views, respectively. Scale bars = 0.2 mm.

Fig. 15. Tylopus acuminatus sp. nov., paratype, Ƌ, from Phufa (A–B), paratype, Ƌ, from Phuthalang forest protected area (C–D), paratype, Ƌ, from forest near road (E), paratype, Ƌ, from Ban Na Thong (F), paratype, Ƌ, from Phagneung Phoukulom waterfall (G–H). A–H. Distal part, sublateral, submesal, dorsal, subdorsal, dorsal, subdorsal, ventral and dorsal views, respectively. Scale bars = 0.2 mm. Red colour shows process h. Green colour shows spine z.

Fig. 25. Tylopus baenzigeri Golovatch & Enghoff, 1993, Ƌ, right gonopod. A–B. Mesal and lateral views, respectively. C–F. Distal part, submesal, sublateral, subcaudal and suboral views, respectively. Scale bar = 0.2 mm.

Fig. 19. Tylopus thunghaihin sp. nov., holotype, Ƌ. A–B. Right gonopod, lateral and mesal views, respectively. C. Leg of segment 9. Scale bars: A–B = 0.2 mm; C = 1 mm. Abbreviations are explained in the text.

Fig. 26. Distribution of the new eight species of Tylopus. Filled triangle. T. punctus sp. nov. Filled circle. T. acuminatus sp. nov. Crossed circle. T. acuminatus sp. nov., T. dorsalis sp. nov. Crossed square. T. acuminatus sp. nov. T. retusus sp. nov. Open circle. T. retusus sp. nov. Open triangle. T. baenzigeri Golovatch & Enghoff, 1993. Filled square. T. hongkhraiensis sp. nov. Open diamond. T. thunghaihin sp. nov. Inverted filled triangle. T. flavolineatus sp. nov. Open square. T. moniliformis sp. nov.

Fig. 4. Tylopus hongkhraiensis sp. nov., holotype, Ƌ (B–J), paratypes, Ƌ, ♀ (A). A. Habitus, live coloration. B–C. Anterior part of body, dorsal and lateral views, respectively. D–E. Segments 10 and 11, dorsal and lateral views, respectively. F–H. Posterior part of body, dorsal, ventral and lateral views, respectively I–J. Sternal cones between coxae 4, subcaudal and sublateral views, respectively. Abbreviations are explained in the text.

Fig. 16. Tylopus dorsalis sp. nov., holotype, Ƌ. A. Habitus, live coloration. B–C. Anterior part of body, dorsal and lateral views, respectively. D–E. Segments 10 and 11, dorsal and lateral views, respectively. F–H. Posterior part of body, dorsal, ventral and lateral views, respectively. I–J. Sternal cones between coxae 4, caudal and sublateral views, respectively.

Fig. 5. Tylopus hongkhraiensis sp. nov., holotype, Ƌ, right gonopod. A–C. Lateral, mesal and suboral views, respectively. Scale bar = 0.2 mm. Abbreviations are explained in the text.

Fig. 6. Tylopus hongkhraiensis sp. nov., holotype, Ƌ, left gonopod. A–B. Mesal and lateral views, respectively. C–F. Distal part, sublateral, submesal, suboral and subcaudal views, respectively. Scale bars = 0.1 mm.

Fig. 8. Tylopus moniliformis sp. nov., paratype, Ƌ, left gonopod. A–B. Mesal and lateral views, respectively. Scale bar = 0.2 mm. Abbreviations are explained in the text.

Fig. 9. Tylopus moniliformis sp. nov., holotype, Ƌ, left gonopod. A–B. Lateral and mesal views, respectively. C–F. Distal part, lateral, mesal, subcaudal and suboral views, respectively. Scale bars = 0.1 mm.

Fig. 22. Tylopus punctus sp. nov., holotype, Ƌ. A–D. Left gonopod, lateral, mesal, subcaudal and suboral views, respectively. Scale bar = 0.2 mm. Abbreviations are explained in the text.