Therates, Latreille, 1817, Latreille, 1817

Wiesner, Jürgen, 2013, The chennelli group of the Genus Therates Latreille (Coleoptera: Cicindelidae) 114. Contribution towards the knowledge of Cicindelidae, Insecta Mundi 2013 (315), pp. 1-86 : 2-9

chennelli group

chennelli group Wiesner 1988: 11 - 15 ; 1992: 89; Probst and Wiesner 1994: 32, 1994a: 100, 101; Sawada and Wiesner 1997: 81 - 83, 2004: 262, 263.

Nomenclatural note. The alternative spelling with one n used by several workers is incorrect. The name should properly be spelled with two n ’s, as established by Bates (1878: 335).

Characters. For the adult specimens of the chennelli group a standard description protocol was followed ( Figure 1 ): body length, excluding labrum (4.8 mm- 11.2 mm); general ground color shining black or greenish black or violet black, pronotum and parts of head in some species reddish. Mandibles yellowish, brownish distally in females, teeth brownish marginally. Labrum mostly as wide as long, yellowish, with dark margin or blackish, lacking basal tooth, with five to seven apical teeth and one lateral tooth. Labial and maxillary palpi completely yellowish or darker distally. Antennae long, medium size or short, slender or lanceolate, distal two antennomeres of male enlarged in some species, scape with a single apical bristle, antennomeres 2 to 5 glabrous, antennomeres 6 to 11 finely and evenly pubescent; scape yellowish above, black on underside, all the other antennomeres brownish or blackish. Clypeus glabrous. Frons smooth with two shallow bumps or a furrow in the posterior part of the orbital plates. Eyes large, prominent. Pronotum glabrous and rounded, generally as wide as long, constricted in front and at back, transverse furrows strong, middle line and lateral lines nearly obsolete, middle line often with several transverse short branches. Scutellum triangular and obvious. Flight wings always present. Elytra with basal and apical hump. distinctly punctate in frons, shallower or absent in the apical part. Elytral apex with or without rounded or angular lateral corner and with angular sutural corner or sutural tooth, recurved or straight between them. Maculations bright and variable in size and shape. including a short, long or broken humeral lunule, a basal dot, that is sometimes absent, a central dot of characteristic shape (e. g. subsquare, fascia like, acutely angled inwards toward the suture, acutely angled outwards toward the front), and an apical dot that may be absent or with an apex that is transparent brownish or yellowish. A central band, if present, is not oriented outward at its anterior end. All or some maculae may connect with others. Mesoepisterna with or without grooves and pits. Elytral epipleura distinct. Ventral body brownish or black, ventrites dark, sometimes brownish marginally. Legs yellowish, femora, tibiae and tarsomeres sometimes darker distally. Legs of females darker as a whole in some species, Metatibiae of females bicolored in some species. Femora nearly glabrous; tibiae sparsely setose, tarsi somewhat more thickly setose; tarsomeres 1 to 4 of males are slightly widened and thickly setose. Length of the aedeagus 1.3 to 2.3 mm, shape straight or curved, with various distinct shaped points distally. Larval forms of the chennelli group are undescribed.

Natural History. Knowledge about the natural history of these species is limited, however, it appears that most, if not all, of these species are found exclusively within the forest in shady and moist areas ( Naviaux and Pinratana 2004: 62). Adults occur primarily on leaves of undergrowth shrubs, much like species of Collyris , watching for their prey. They are extremely shy and their flight is powerful. Sometimes, after flying away, they return to land on the same leaf.

Key to the Therates species groups (based on Wiesner 1988).

1. Fourth metatasomere lacking setigerous bottom, instead with rigid and erected bristles distally. ......................................................................................................................... spectabilis group

— Fourth metatarsiomere with setigerous bottom ........................................................................... 2

2(1). Clypeus with two erect setae ......................................................................................................... 3

— Clypeus glabrous ............................................................................................................................ 4

3(2). Frons entirely curvate ........................................................................................... festivus group

— Frons geniculate in front ..................................................................................... labiatus group

4(2). Elytra containing apical hump only .................................................................. cribratus group

— Elytra with more than one hump ................................................................................................. 5

5(4). Elytra with basal, central and apical hump ........................................................... batesii group

— Elytra with basal and apical hump only ....................................................................................... 6

6(5). Labrum with basal tooth ............................................................................................................... 9

— Labrum lacking basal tooth .......................................................................................................... 7

7(6). Elytra with bright central band, oriented outward at its anterior end ............ obliquus group

— Elytra with bright central dot or transverse rectangular central band ..................................... 8

8(7). Palpi bright ......................................................................................................... chennelli group

— Distal segment of labial palpi and maxillary palpi as a whole dark ................ tuberosus group

9(6). Elytra with bright central band, oriented outward at its anterior end ............ obliquus group

— Maculation of elytra not as above ............................................................................................... 10

10(9). Basal three metatarsomeres cylindrically enlarged ............................................. hennigi group

— Metatarsi not as above ................................................................................................................ 11

11(10). Elytral apex rounded or elytral apex drawn backwards or pronotum more constricted in front than at back .................................................................................................... coeruleus group

— Elytral apex not rounded or elytral apex not drawn backwards or pronotum more constricted at back than in front or equally in front and at back ................................................................. 12

12(11). Elytral apex with produced sutural spine or prominent triangular sutural angle; pronotum as a rule similarly constricted in front and at back .......................................... spinipennis group

— Elytral apex without produced sutural spine; pronotum more constricted at back than in front ........................................................................................................................... fasciatus group

Remarks. The specimens studied were also analyzed using numerical cladistics ( Camin and Sokal 1965). Figure 2 lists the characters used together with the matrix of OTU’s (operational taxonomic units). Figure 3 shows the provisional tree resulting from the cladistic analysis of the species. I tentatively use the order, in which the species appear in the tree, to arrange them within this paper.

List of the species of chennelli group

One species is known from Nepal, eight from India, seven from Myanmar, 11 from Thailand, three from Malaysia, 21 from Laos, 13 from Vietnam, and three from China. At the time of its description ( Werner 1991: 230), Therates hiermeieri was included in chennelli group, but later placed it close to the Th. fleutiauxi Horn, 1898 by Sawada and Wiesner (2003: 50). Th. hiermeieri , due to its basal tooth on the labrum, will key out in the identification key to Therates groups as belonging to coeruleus group.

1. montaneus Werner, 1992 China.

2. moraveci Sawada and Wiesner, 1999 Laos.

3. pseudorugifer pseudorugifer Sawada and Wiesner, 1999 Laos.

3a. pseudorugifer pentalabiodentatus Matalin, 2001 China.

4. baolocensis Wiesner, 1996 , new rank Vietnam.

5. apiceflavus Sawada and Wiesner, 1999 Laos.

6. laotiensis Sawada and Wiesner, 1999 Laos.

7. topali Mandl, 1972 Vietnam.

8. bannokcolus Sawada and Wiesner, 1999 Laos.

9. circumscriptus Moravec and Wiesner, 1999 Laos.

10. probsti Wiesner, 1988 Laos, Vietnam.

11. vietnamensis Wiesner, 1988 Vietnam.

12. concinnus Gestro, 1888 Myanmar, Thailand, Laos, Vietnam. 13. bannapecolus Sawada and Wiesner, 1999 Laos.

14. major Probst and Wiesner, 1994 Thailand.

15. annandalei Horn, 1908 India.

16. nepalensis Probst and Wiesner, 1994 Nepal, India.

17. gestroi Horn, 1900 Laos, Vietnam.

18. dembickyi Sawada and Wiesner, 2002 Laos.

19. kraatzi Horn, 1900 Thailand, Malaysia.

20. similis Probst and Wiesner, 1994 Thailand.

21. pearsoni new species Vietnam.

22. myanmarensis Wiesner, 1999 Myanmar.

23. confluens Wiesner, 1988 Malaysia, Vietnam.

24. pacholatkoi Sawada and Wiesner, 2004 Laos.

25. pseudoconfluens Sawada and Wiesner, 1999 Laos, China.

26. pseudochenelli pseudochenelli Probst and Wiesner, 1994 Thailand, Malaysia.

26a. pseudochenelli rufus Probst and Wiesner, 1994 Thailand.

27. safraneki new species Laos.

28. nagaii Sawada and Wiesner, 2000 Myanmar.

29. schuelei new species Vietnam.

30. rugifer Horn, 1902 Vietnam.

31. phongsalyensis Sawada and Wiesner, 2004 Laos.

32. namthacolus Sawada and Wiesner, 1999 Laos.

33. sigridgeissleri new species Laos.

34. tonkinensis Horn, 1902 Vietnam.

35. jendeki Sawada and Wiesner, 1997 India.

36. csorbai Wiesner, 1999 Laos.

37. apicenigrus Sawada and Wiesner, 1999 Laos.

38. rihai Moravec and Wiesner, 2001 Thailand.

39. chennelli Bates, 1878 Myanmar.

40. pseudoprobsti Probst and Wiesner, 1994 Thailand.

41. arunachalcolus Sawada and Wiesner, 2006 India.

42. ingridae Sawada and Wiesner, 2006 India.

43. nigromarginalis Probst and Wiesner, 1994 Myanmar, Thailand.

44. murzini Wiesner, 1999 Myanmar.

45. haucki Moravec and Wiesner, 2001 Thailand, Laos, Vietnam.

46. sausai Sawada and Wiesner, 1997 India.

47. miyamai Sawada and Wiesner, 2000 Myanmar.

48. dohertyi Horn, 1905 India.

49. ottomerkli Wiesner, 1999 Laos.

50. clavicornis Horn, 1902 Vietnam.

51. rogeri Probst and Wiesner, 1994 Thailand.

52. khaoyaii new species Thailand.

53. waagenorum Horn, 1900 India.

Key to the members of chennelli group

In cases of doubt, comparision of the aedeagus with the figures in this paper may be helpful.

1. Ventral aspect completely pale; metaepisternum yellow to reddish brown ................................. 2

— Ventral aspect partly to almost dark; metaepisternum black ...................................................... 7

2(1). Elytral maculation including the middle suture from base to apex (6.4 mm, Thailand) .............. ......................................................................................... 51. Th. rogeri Probst and Wiesner

— Elytral maculation not as above ................................................................................................... 3

3(2). Pronotum brownish ....................................................................................................................... 4

— Pronotum black ............................................................................................................................. 5

4(3). Male with distal two antennomeres obviously elongated (6.2 mm- 7.4 mm, Vietnam) .................. ........................................................................................................... 50. Th. clavicornis Horn

— Male without distal antennomeres elongated (6.4 mm- 6.6 mm, Laos) .......................................... ....................................................................................................... 49. Th. ottomerkli Wiesner

5(3). Central dot forming a narrow band (6.3 mm- 7.6 mm, India) ................ 48. Th. dohertyi Horn

— Central dot not as above ................................................................................................................ 6

6(5). Distance between elytral apical hump and lateral tooth short; aedeagus short, less prominent (5.8 mm- 7.5 mm, Laos) ................................................ 28. Th. nagaii Sawada and Wiesner

— Distance between elytral apical hump and lateral tooth long; aedeagus long and prominent (6.5 mm- 7.4 mm, Myanmar) .................................................................... 44. Th. murzini Wiesner

7(1). Frons mostly reddish ..................................................................................................................... 8

— Frons mostly black ........................................................................................................................ 9

8(7). Elytral punctures isolated (5.7 mm- 6.2 mm, India) ....................... 53. Th. waagenorum Horn

— Elytral punctures connected in short rows (6.6 mm-8.0 mm, Thailand) ....................................... .................................................................................................. 52. Th. khaoyaii new species

9(7). Elytral maculae variably connected throughout ........................................................................ 10

— Elytral maculae isolated, or at most humeral lunule and basal dot connected at base and/or humeral lunule connected with central dot ............................................................................ 22

10(9). Labrum entirely bright yellow .................................................................................................... 12

— Labrum dark at outer margin ..................................................................................................... 11

11(10). Labrum with complete broad black outer margin (6.0 mm-7.0 mm, Myanmar, Thailand) .......... ..................................................................... 43. Th. nigromarginalis Probst and Wiesner

— Labrum with incomplete blackish outer margin from lateral tooth up to base (6.2 mm- 8.1 mm, India) ............................................................................. 46. Th. sausai Sawada and Wiesner

12(10). Dark wedge shaped recession between humeral lunule and central dot absent or small ........ 13

— Dark wedge shaped recession between humeral lunule and central dot obvious and extends from marginal suture to center ........................................................................................................ 17

13(12). Ventrites black or with narrow yellow lateral margin (6.8 mm- 7.5 mm, Malaysia, Vietnam) ..... ........................................................................................................ 23. Th. confluens Wiesner

— Ventrites with broad and distinctive lateral yellow margins ..................................................... 14

14(13). Basal hump without black spot .................................................................................................. 16

— Basal hump with black spot ........................................................................................................ 15

15(14). Humeral lunule and central dot not extending to the middle suture (7.3 mm- 8.2 mm, Laos) ..... ................................................................................................. 27. Th. safraneki new species

— Humeral lunule and central dot extending to the middle suture (6.3 mm- 8.3 mm, Laos, China) .................................................................... 25. Th. pseudoconfluens Sawada and Wiesner

16(14). Apical dot extended forward medially (6.3 mm-8.0 mm, Thailand, Laos, Vietnam) ...................... .................................................................................... 45. Th. haucki Moravec and Wiesner

— Apical dot not extended forward medially (5.8 mm- 7.6 mm, Laos) ................................................ ............................................................................. 24. Th. pacholatkoi Sawada and Wiesner

17(12). Posterior margin of central dot angled gradually outwards toward the front ......................... 19

— Posterior margin of central dot nearly right-angled to elytral edge ......................................... 18

18(17). Humeral lunule and central dot almost completely connected (7.5 mm- 8.5 mm, Myanmar) ....... ............................................................................................................. 39. Th. chennelli Bates

— Humeral lunule and central dot almost isolated from each other (7.2 mm- 9.1 mm, Myanmar, Thailand, Laos, Vietnam) .............................................................. 12. Th. concinnus Gestro

19(17). Central dot acutely angled outwards toward the front .............................................................. 20

— Central dot not as above; aedeagus with roundish tip .............................................................. 21

20(19). Ventrites with narrow yellowish lateral margin; aedeagus with club-shaped tip (6.5 mm- 7.6 mm, Thailand) ........................................................................ 20. Th. similis Probst and Wiesner

— Ventrites with broad yellowish lateral margin; aedeagus with evenly pointed tip (6.8 mm- 8.2 mm, Thailand) ...................................................................... 38. Th. rihai Moravec and Wiesner

21(19) Elytral maculation yellowish (7.0 mm- 8.7 mm, Thailand, Malaysia) ............................................ ............................................ 26. Th. pseudochenelli pseudochenelli Probst and Wiesner

— Elytral maculation red brown (7.5 mm- 8.6 mm, Thailand) ............................................................ ............................................................ 26a. Th. pseudochenelli rufus Probst and Wiesner

22(9). Humeral lunule short, not extended onto disk .......................................................................... 23

— Humeral lunule long, extended onto disk and diverging away from marginal suture ............. 29

23(22). Central dot subsquare or trapezoidal ......................................................................................... 24

— Central dot not as above .............................................................................................................. 27

24(23). Antennae short, male with distal two antennomeres obviously enlarged (8.0 mm- 11.2 mm, Vietnam) .......................................................................................................... 34. Th. tonkinensis Horn

— Antennae longer, male without distal two antennomeres enlarged .......................................... 25

25(24). Labrum dark at outer margin (6.5 mm- 7.5 mm, Laos) .................................................................. ....................................................................... 31. Th. phongsalyensis Sawada and Wiesner

— Labrum uniformly pale ................................................................................................................ 26

26(25). Labrum with six apical teeth; body size smaller (8.2 mm- 8.7 mm, Laos) ...................................... ................................................ 3. Th. pseudorugifer pseudorugifer Sawada and Wiesner

— Labrum with five apical teeth; body size larger (9.5 mm, China) .................................................. ........................................................... 3a. Th. pseudorugifer pentalabiodentatus Matalin

27(23). Labrum dark at outer margin ..................................................................................................... 28

— Labrum uniformly pale (7.4 mm, Laos) ........... 9. Th. circumscriptus Moravec and Wiesner

28(27). Male with distal two antennomeres obviously elongated; body size usually less than 8 mm (6.7 mm- 8.2 mm, Vietnam) ............................................................................. 30. Th. rugifer Horn

— Male without distal two antennomeres elongated; body size usually greater than 8 mm (8.0 mm-9.2 mm, Laos) ................................................... 32. Th. namthacolus Sawada and Wiesner

29(22). Central dot subsquare or a trapezoidal band ............................................................................. 30

— Central dot more or less roundish, not forming a band ............................................................ 34

30(29). Labrum uniformly yellow ............................................................................................................ 31

— Labrum dark at outer margin (7.8 mm, India) ............................................................................... ..................................................................... 41. Th. arunachalcolus Sawada and Wiesner

31(30). Humeral lunule slender (6.9 mm- 8.2 mm, China) ............................ 1. Th. montaneus Werner

— Humeral lunule broad ................................................................................................................. 32

32(31). Yellow color of elytral apex reaches the apical humps (7.7 mm- 8.2 mm, Nepal, India) ................ ................................................................................. 16. Th. nepalensis Probst and Wiesner

— Yellow apical dot does not reach the apical humps .................................................................... 33

33(32). Humeral lunule long, tongue like and enlarged towards central dot (8.2 mm- 9.6 mm, Laos) ..... ......................................................................... 13. Th. bannapecolus Sawada and Wiesner

— Humeral lunule shorter and uniform in width throughout (9.0 mm, Thailand) .......................... .......................................................................................... 14. Th. major Probst and Wiesner

34(29). Central dot acutely angled outwards toward the front .............................................................. 35

— Central dot not as above .............................................................................................................. 45

35(34). Elytral apex with a yellow dot .................................................................................................... 42

— Elytral apex with no yellow dot .................................................................................................. 36

36(35). Labrum uniformly pale ................................................................................................................ 38

— Labrum dark at outer margin ..................................................................................................... 37

37(36). Ventrites black to margins (6.7 mm- 7.6 mm, Vietnam) .............. 29. Th. schuelei new species

— Ventrites with brownish margins (4.8 mm- 6.7 mm, India) ............................................................. .................................................................................. 42. Th. ingridae Sawada and Wiesner

38(36). Male with antennae short, reaching elytral shoulders (5.7 mm- 6.9 mm, Laos) ........................... ............................................................................. 8. Th. bannokcolus Sawada and Wiesner

— Male with antennae longer, reaching behind elytral shoulders ................................................ 39

39(38). Apex of elytra black ..................................................................................................................... 40

— Apex of elytra transparent brownish or yellowish ..................................................................... 41

40(39). Elytra without basal dot (7.0 mm- 7.4 mm, Laos) ........................................................................... ............................................................................. 37. Th. apicenigrus Sawada and Wiesner

— Elytra with basal dot (6.9 mm- 7.5 mm, Laos) ............. 2. Th. moraveci Sawada and Wiesner

41(39). Central dot horizontal (6.8 mm- 7.1 mm, Laos) ....... 18. Th. dembickyi Sawada and Wiesner

— Central dot acutely angled outwards toward the front (5.4 mm- 7.2 mm, Laos, Vietnam) ........... ................................................................................................................... 17. Th. gestroi Horn

42(35). Humeral lunule short .................................................................................................................. 43

— Humeral lunule long, nearly reaching the central dot .............................................................. 44

43(42). Labrum uniformly yellowish (7.3 mm- 8.8 mm, Laos, Vietnam) .......... 10. Th. probsti Wiesner

— Labrum brownish laterally (6.9 mm- 7.5 mm, Laos) ........ 33. Th. sigridgeissleri new species

44(42). Ventrites black to margins (7.5 mm- 8.4 mm, Vietnam) ................. 4. Th. baolocensis Wiesner

— Ventrites with yellowish margins (6.5 mm- 7.5 mm, Laos) .................. 36. Th. csorbai Wiesner

45(34). Humeral lunule long, connected with the central dot or nearly reaching it ............................ 46

— Humeral lunule shorter ............................................................................................................... 49

46(45). Humeral lunule broad and strongly extended onto disk ........................................................... 47

— Humeral lunule narrow and barely extending onto disk ........................................................... 48

47(46). Light color of elytral apex reaching but not extending onto apical humps (6.7 mm- 8.4 mm, Thailand, Malaysia) ............................................................................... 19. Th. kraatzi Horn

— Light color of elytral apex covering apical humps (7.2 mm- 8.2 mm, India) .................................. ......................................................................................................... 15. Th. annandalei Horn

48(46). Apex of elytra broadly yellow (6.1 mm- 7.8 mm, Laos) .................................................................... ................................................................................ 5. Th. apiceflavus Sawada and Wiesner

— Apex of elytra yellow only at a small area near the suture (5.8 mm- 7.7 mm, Vietnam) ............... .................................................................................................................... 7. Th. topali Mandl

49(45). Elytral apex black with large and bright apical dot ................................................................... 50

— Elytra with small, yellow apical dot at suture or apex transparent dark brown ..................... 51

50(49). Central elytral dot broad; male without distal two antennomeres elongated (6.4 mm- 9.1 mm, Laos) ......................................................................... 6. Th. laotiensis Sawada and Wiesner

— Central elytral dot slender; male with distal two antennomeres obiously elongated (7.3 mm- 8.6 mm, Myanmar) ..................................................................... 22. Th. myanmarensis Wiesner

51(49). Basal elytral dot longish and located near middle suture ......................................................... 52

— Basal elytral dot subsquare roundish and located besides the scutellum ................................. 54

52(51). Basal elytral dot broad; male with distal two antennomeres obiously elongated (8.2 mm- 9.2 mm, Vietnam) .................................................................................... 21. Th. pearsoni new species

— Basal elytral dot narrow, male without distal two antennomeres elongated ............................ 53

53(52). Central elytral dot broad (7.2 mm- 8.7 mm, Thailand) .................................................................... ........................................................................... 40. Th. pseudoprobsti Probst and Wiesner

— Central elytral dot slender (5.8 mm- 7.3 mm, Myanmar) ................................................................ ................................................................................... 47. Th. miyamai Sawada and Wiesner

54(51). Central elytral dot subsquare; aedeagus with long pointed tip, knobbed apically (6.7 mm- 9.1 mm, India) ........................................................................... 35. Th. jendeki Sawada and Wiesner

— Central dot acutely angled inwards toward the suture; aedeagus with thick tip (6.3 mm- 8.7 mm, Vietnam) ................................................................................... 11. Th. vietnamensis Wiesner

Figures

Figure 1. Therates spec., left = ventral view, right = dorsal view: (1) mandible, (2) labrum, (3) clypeus, (4) eye, (5) orbital plate, (6) pronotum, (7) lateral line, (8) middle line, (9) scutellum, (10) basal hump, (11) apical hump, (12) apical dot, (13) central dot, (14) humeral lunule, (15) basal dot, (16) transverse furrows, (17) antenna, (18) frons, (19) maxillary palpus, (20) labial palpus, (21) mesoepisternum, (22) epipleurum, (23) ventrites. Scale = 1 mm.

Figures 3. Provisional arrangement of species of the chennelli group following the results of cladistic analysis.