Hyleoglomeris Verhoeff, 1910

Golovatch, Sergei I., Geoffroy, Jean-Jacques & Mauriès, Jean-Paul, 2006, Review of the millipede genus Hyleoglomeris Verhoeff, 1910 (Diplopoda, Glomerida, Glomeridae), with descriptions of new species from caves in Southeast Asia, Zoosystema 28 (4), pp. 887-915 : 889-894

Genus Hyleoglomeris Verhoeff, 1910

Hyleoglomeris Verhoeff, 1910: 245 , figs 1-4.

Nesoglomeris Carl, 1912: 100 . Type species: Nesoglomeris sarasinorum Carl, 1912 , by subsequent designation ( Jeekel 1970: 15); synonymised by Verhoeff (1912: 150).

Sundameris Verhoeff, 1936: 163 . Type species: Apiomeris jacobsoni Silvestri, 1917 , by monotypy; synonymised by Hoffman (1980: 68).

Perkeomeris Verhoeff, 1936: 163 (invalidly proposed, without type species designation).

Okeanomeris Verhoeff, 1942: 214, figs 18, 19. Type species: Hyleoglomeris nigra Verhoeff, 1942 , by monotypy; synonymised by Hoffman (1980: 68).

Zygethomeris Chamberlin, 1921: 55 . Type species: Zygethomeris lamprus Chamberlin, 1921 , by original designation; synonymised by Hoffman (1980: 68).

NOTE. — Since Mauriès (1971) referred to Sundameris and Okeanomeris as “synonyms or subgenera” of Hyleoglomeris , we emphasize that it was Hoffman (1980) who first listed them as strict synonyms.

TYPE SPECIES. — Hyleoglomeris multilineata Verhoeff, 1910 , by subsequent designation ( Silvestri 1917: 107).

SPECIES INCLUDED

Below is a checklist of the species currently attributed

to Hyleoglomeris , arranged in alphabetical order.

1. H. albicollis Golovatch, 1983 , described from Thailand (Ghi NP, Mae Chaem), still known only from the original description ( Golovatch 1983).

2. H. albicornis ( Pocock, 1894) , originally described from Sumatra (Singkarah) as Glomeris albicornis Pocock, 1894 , cited as such also by Silvestri (1895), still known only from the original description ( Pocock 1894).

3. H. albicorporis Zhang & Zhang, 1995 , described rather recently from a cave at Baoshan, Yunnan Province, China ( Zhang & Zhang 1995).

4. H. alticola ( Carl, 1912) , first described as Nesoglomeris alticola Carl, 1912 , from Bowonglangi, S Sulawesi; still known only from the original description ( Carl 1912).

5. H. armeniaca Golovatch, 1989 , described and still known only from N Armenia, Caucasus ( Golovatch 1989).

6. H. atricornis ( Silvestri, 1917) , first described as Apiomeris (Hyleoglomeris) atricornis Silvestri, 1917 , from Borneo (Mt. Mulu); still known only from the original description ( Silvestri 1917).

7. H. awchasica (Brandt, 1840) , from Colchis, W Caucasus within Georgia and Russia, with both Glomeris kallipygos Attems, 1907 and Glomeris kubana Verhoeff, 1921 established as its junior synonyms ( Golovatch 1989).

8. H. beccarii ( Silvestri, 1917) , first described as Apiomeris (Hyleoglomeris) beccarii Silvestri, 1917 , from Sumatra (Mt. Singalan), still known only from the original description ( Silvestri 1917).

9. H. beroni Mauriès, 1984 , described and still known only from Greece (Naxos Island:Zeus Cave) ( Mauriès 1984).

10. H. crassipes Golovatch, 1987 , described and still known only from Nepal (Terhathum District,Tinjura Dara near Chauki) ( Golovatch 1987).

11. H. crebristriata ( Silvestri, 1917) , first described as Apiomeris (Hyleoglomeris) crebristriata Silvestri, 1917 , from Sarawak, Borneo; still known only from the original description from female material ( Silvestri 1917). 12. H.cremea Golovatch, 1983 , described and still known only from N Thailand (Chiang Dao) ( Golovatch 1983). 13. H. diversicolor ( Silvestri, 1895) , first described as Glomeris diversicolor Silvestri, 1895 , from Sumatra (Si-Rambé) ( Silvestri 1895), then redescribed as Apiomeris (Hyleoglomeris) diversicolor ( Silvestri, 1895) , based on type material ( Silvestri 1917). 14. H. electa ( Silvestri, 1917) , first described from female material as Apiomeris (Hyleoglomeris) electa Silvestri, 1917 , from India (Darjeeling District: Ghumti) ( Silvestri 1917), later redescribed based on male and female samples from near Dalat, Vietnam ( Attems 1938), also reported from Xieng Kuang, Boloven Plateau, Laos ( Attems 1953). Some additional comments concerning the identity of this species are given below. 15. H. emarginata Golovatch, 1981 , described from China (Kiangsu Province: Cisian-Shan 25 km S of Nanjing) ( Golovatch 1981), it has since been reported from S Korea as well ( Mikhaljova & Lim 2000); redescribed below. 16. H. epirotica ( Mauriès, 1966) , first described as Speleoglomeris (sic) epirotica Mauriès, 1966 , from Greece (Epirus, Joànnina, Pérama: Pérama Cave) ( Mauriès 1966), transferred to Hyleoglomeris by Mauriès (1984). 17. H. eremita ( Carl, 1912) , first described as Nesoglomeris eremita Carl, 1912 , from a single female holotype from Bowonglangi, S Sulawesi; still known only from the original description ( Carl 1912). 18. H. formosa ( Silvestri, 1895) , first described as Glomeris formosa Silvestri, 1895 , from Sumatra (Benkoelen) ( Silvestri 1895), later redescribed as Apiomeris (Hyleoglomeris) formosa ( Silvestri, 1895) , based on type material ( Silvestri 1917). 19. H. gorkhalis Golovatch, 1987 , described and still known only from Nepal (Gorkha District, between Naya Sangu and Gorkha) ( Golovatch 1987). 20. H. insularum Verhoeff, 1936 , first described as Hyleoglomeris (Perkeromeris) insularum Verhoeff, 1936 , based on a single female holotype from near Tokyo, Japan ( Verhoeff 1936); later again reported from the same locality ( Chamberlin & Wang 1953; Shinohara 1981). 21. H. jacobsoni ( Silvestri, 1917) , first described as Apiomeris (Hyleoglomeris) jacobsoni Silvestri, 1917 , from Java (Nongkodjadjar) ( Silvestri 1917), later reported from several Lesser Sunda islands: Bali (Gitgit), Sumbawa (Batoe Doelang) and Flores (Geli Moetoe) ( Attems 1930). Based solely on 3- rather

than 4-segmented male telopodites 17 as illustrated by Attems (1930), Verhoeff (1936) established Sundameris to only incorporate H. jacobsoni . The conspecificity of Attems’ samples with type material is still to be verified (cf. Verhoeff 1936).

22. H. japonica Verhoeff, 1936 , first described as Hyleoglomeris (Perkeromeris) japonica Verhoeff, 1936 , from Enoshima near Tokyo, Japan ( Verhoeff 1936).

23. H. khumbua Golovatch, 1987 , described and still known only from Nepal (Solukhumbu District, Khumbu, Mt. Everest region) ( Golovatch 1987).

24. H. kirgisica Golovatch, 1976 , described and still known only from the W Tian-Shan Mountains within Kirghizia, Central Asia ( Golovatch 1976; Read & Golovatch 1994).

25. H. kirropeza ( Attems, 1897) , first described as Glomeris kirropeza Attems, 1897 , from Sulawesi (Minahassa) from a few females ( Attems 1897), later redescribed as Nesoglomeris kirropeza ( Attems, 1897) from both sexes from Lokon and Soputan, N Sulawesi ( Carl 1912).

26. H. koreana Golovatch, 1978 , first described from S Korea ( Golovatch 1978), currently known over most of the Korean Peninsula and on Cheju Island ( Golovatch 1981; Lim et al. 1992; Mikhaljova & Kim 1993; Kim & Lim 1995a, b; Mikhaljova & Lim 2000; Mikhaljova et al. 2000).

27. H. lamprus ( Chamberlin, 1921) , first described as Zygethomeris lamprus Chamberlin, 1921 , from Borneo ( Sarawak, Ladong) ( Chamberlin 1921), it has since been recorded in the Philippines (Samar) ( Wang 1951, 1961). However, like most of Wang’s work, this identification requires verification.

28. H. lenkorana Golovatch, 1976 , described and still known from Hyrcania within Azerbaijan, Caucasus and N Iran ( Golovatch 1976, 1989).

29. H. lohmanderi Golovatch, 1975 , described and still known only from Azerbaijan, Caucasus ( Golovatch 1975, 1989).

30. H. lucida Haga, 1956 , described and since referred to as Hyleoglomeris lucidus Haga, 1956 (recte: lucida , to agree in gender with the feminine generic name), from Honshu, Japan ( Takashima & Haga 1956; Shinohara 1978).

31. H. maior Attems, 1938 , described and still known only from two localities in S Vietnam: Phanrang and Hon Ba ( Attems 1938). Some additional comments concerning the identity of this species are given below.

32. H. minuta Verhoeff, 1910 , described from Borneo ( Verhoeff 1910), redescribed based on type material from Mt. Radjang, Klawang, Borneo ( Silvestri 1917).

33. H. modesta ( Silvestri, 1917) , described as Apiomeris (Hyleoglomeris) modesta Silvestri, 1917 , from India (Assam, Kobo); still known only from the original description ( Silvestri 1917).

34. H. modiglianii ( Silvestri, 1895) , described as Glomeris Modiglianii (sic) Silvestri, 1895, from Nias Island (Lelemboli) ( Silvestri 1895), redescribed as Apiomeris (Hyleoglomeris) modiglianii ( Silvestri, 1917) , based on type material ( Silvestri 1917).

35. H. montana Golovatch, 1983 , described and still known only from N Thailand (Chieng Mai, Doi Inthanon) ( Golovatch 1983).

36. H. multilineata Verhoeff, 1910 , described from Borneo (Bengkajong) ( Verhoeff 1910, 1915), redescribed from type material ( Silvestri 1917).

37. H. nagarjunga Golovatch, 1987 , described and still known only from Nepal (Kathmandu Valley, Nagarjung, Mt. Jamacok) ( Golovatch 1987).

38. H. nigra Verhoeff, 1942 , described as Hyleoglomeris (Okeanomeris) nigra Verhoeff, 1942 , from Japan (Shikoku, near Koti) ( Verhoeff 1942).

39. H. paucilineata ( Silvestri, 1917) , described as Apiomeris (Hyleoglomeris) paucilineata Silvestri, 1917 , based solely on female material from Borneo (Kari Orang, Kutei); still known only from the original description ( Silvestri 1917).

40. H. piccola ( Attems, 1899) , described and securely known only from Hyrcania within Azerbaijan, Caucasus ( Attems 1899; Golovatch 1976, 1989).

41. H. pulchra Attems, 1953 , described based solely on female material, still known only from Laos (Paklay) ( Attems 1953).

42. H. robusta Attems, 1938 , described based solely on female material from Peak Lang Biang, S Vietnam; hitherto known only from the original description ( Attems 1938), redescribed and commented upon below.

43. H. sakamotoensis Takano, 1981 , described and still known only from Kyushu, Japan ( Takano 1981).

44. H.sarasinorum ( Carl, 1912) , described as Nesoglomeris sarasinorum Carl, 1912 , from S Sulawesi (Loka); still known only from the original description ( Carl 1912).

45. H. siamensis ( Silvestri, 1917) , described as Apiomeris (Hyleoglomeris) siamensis Silvestri, 1917 , from Thailand (Meetaw, Raheng); still known only from the original description ( Silvestri 1917).

46. H. sinensis ( Brölemann, 1896) , described as Glomeris sinensis Brölemann, 1896 , from China ( Brölemann 1896); redescribed below.

47. H. specialis Golovatch, 1989 , described and still known only from the Central and E Caucasus Major within Georgia, Azerbaijan and Russia ( Golovatch 1989).

48. H. stuxbergi ( Attems, 1909) , described as Glomeris Stuxbergi (sic) Attems, 1909, from Mangaesi, foot of Mt. Fuji, Honshu, Japan ( Attems 1909), later reported from other localities in Saitama Prefecture, Honshu ( Shinohara 1978).

49. H. sulcata Verhoeff, 1942 , described and still known only from female material taken from near Takashima (Tokushima?), Shikoku, Japan ( Verhoeff 1942); omitted from the later keys to the Japanese Hyleoglomeris species ( Takakuwa 1954; Miyosi 1959).

50. H. talasensis ( Lohmander, 1939) , described as Glomeris talasensis Lohmander, 1939 , from Central Turkey (Kayseri,Talas) ( Lohmander 1939), redescribed from type and additional topotypic material ( Golovatch 1989).

51. H. tinjurana Golovatch, 1987 , described and still known only from Nepal (Terhathum District,Tinjura Dara near Chauki) ( Golovatch 1987).

52. H. triangularis Haga, 1968 , described and still known only from a cave in Fukuoka District, Kyushu, Japan ( Haga 1968).

53. H. triangulifera Attems, 1938 , described and still known only from female material taken from near Nhatrang, S Vietnam ( Attems 1938).

54. H. uenoi Miyosi, 1955 , described and still known only from Yamaguchi Prefecture, S Honshu, Japan ( Miyosi 1955).

55. H. venustula ( Silvestri, 1917) , described as Apiomeris (Hyleoglomeris) venustula Silvestri, 1917 , from India (NE Assam, Sadiya), still known only from its original description ( Silvestri 1917).

56. H. vittata Verhoeff, 1929 , described from Taiwan (Kankan), still known only from its original description ( Verhoeff 1929).

57. H. yamashinai Verhoeff, 1937 , described as Hyleoglomeris (Perkeomeris) yamashinai Verhoeff, 1937 , from Okinawa Island, Riukius, Japan ( Verhoeff 1937), redescribed based on topotypic material ( Murakami 1975).

58. H. zonifera ( Silvestri, 1917) , described as Apiomeris (Hyleoglomeris) zonifera Silvestri, 1917 , based solely on female material from Borneo (Mt. Matang); still known only from the original description ( Silvestri 1917).

In addition, at least one unidentified Hyleoglomeris

species has been recorded in Taiwan ( Korsós 2004),

and another, apparently new species, occurs in

Tajikistan, Central Asia (Golovatch unpubl.). This

brings the total number of Hyleoglomeris species to

nearly 60, undoubtedly with many more awaiting

discovery and description. This statement is here

illustrated by the descriptions of six new species

from China and Indochina.

No attempt has been made yet to key all Hyleo-

glomeris species, though several regional keys are

available. These concern the faunas of Sulawesi ( Carl

1912), Indochina ( Attems 1938; Golovatch 1983),

the Himalayas ( Golovatch 1987), the Caucaso-

Irano-Anatolian region ( Golovatch 1989) and, in part, Japan ( Verhoeff 1936, 1942; Takakuwa 1954; Miyosi 1959; Murakami 1975). The present paper is therefore concluded by a general key to all 64 species of Hyleoglomeris described to date.

Regrettably, the quality of most of the earlier descriptions and illustrations is often quite poor and many species have been based on females alone. All of which severely hampers taxonomic work on Hyleoglomeris . Most of the old species require restudy, preferably based on fresh topotypic material, in order to attempt a serious revision of this genus in a step-by-step way. Thus, three old species are redescribed herein, one based on type material, the others on topotypes. So it is by necessity that the key below is mainly based on regional faunas rather than being purely systematic. It still contains several problems, especially as regards couplets 24 and 35, which cannot be resolved using the available literature alone.

Hyleoglomeris emarginata Golovatch, 1981 (Fig. 2)

MATERIAL EXAMINED. — China. Kiangsu Province , Nanjing, Zijin (Purple) Mt., 350-450 m, 9.X.1988, leg. P. Beron, 36 ♂♂, 32 ♀♀,  juvs ( NMNHS) ; 3 ♂♂, 3 ♀♀ ( MNHN CC160 ) ; 3 ♂♂, 3 ♀♀ ( ZMUM) .

DESCRIPTION

Length of adults of both sexes ranging between 6.0 and 9.0 mm, width between 3.0 and 4.0 mm. Background coloration varying from castaneous or gray-brown to blackish, legs from whitish yellow to red-brown, often becoming darker distally; markings on terga ranging from whitish to gray-yellow.

Head above level of antennal sockets marbled reddish-brown to dark brown, antennae dark redbrown, antennomere 6 about 2 times longer than wide; ocelli 4 + 1 + 1 to 7 + 1, black, convex. Collum dark, usually with a transverse, central, marbled yellow-brown spot.

Body with a distinctive, more or less wide, yellow to grayish, more or less complete, axial stripe usually growing thinner and more obscure toward pygidium (Fig. 2A). In particularly dark specimens, the pattern is marbled gray-brown and indistinct.Thoracic shield

B

C

D

FIG. 2. — Hyleoglomeris emarginata Golovatch, 1981 , ♂ near-topotypes: A, typical colour pattern; B, rather usual colour pattern on thoracic shield; C, same on a midbody tergite; D, same on pygidium; E, leg 17; F, leg 18; G, leg 19 (telopod), front view; H, distal part of telopod, caudal view. Scale bar: A-D, drawn not to scale; E-H, 0.4 mm.

dark, with a conspicuous, more or less wide, yellow to (medially) grayish band along lateral and anterior margins (Fig.2B); sometimes the band even extends caudolaterally (Fig. 2A), often interrupted near axial stripe; latter sometimes incomplete caudally, always flanked by a paramedian pair of large, more or less marbled spots (Fig. 2A, B). Middorsal spots on each of terga 3 to 11 usually rather parallel-sided, but quite often V-shaped, subtriangular; each of these terga also with a pair of large, sublateral, yellow to marbled yellow-brownish spots which normally do not reach the translucid caudal and lateral edges (Fig. 2A, C). Pygidium with a large, subtriangular to drop-shaped central spot, usually, but not always, set off from a translucid caudal margin (Fig. 2A, D).

Collum with two transverse striae.

Thoracic shield with a relatively narrow hyposchism nearly reaching the caudal tergal contour; 6 or 7 transverse striae, of which 1 or 2 start above schism and, together with subsequent 2 or 3 striae, cross the dorsum.

Male pygidium usually slightly sinuate medially at caudal margin (Fig. 2D).

Male leg 17 (Fig. 2E) with a more or less high, often irregularly rounded, outer coxal lobe; telopodite 4-segmented.

Male leg 18 (Fig. 2F) with a more or less ogival syncoxital notch; telopodite 4-segmented.

Telopods (Fig. 2G, H) with a more or less high but invariably distinctly emarginate central syncoxital lobe flanked by two setose horns crowned with a lanceolate structure, with or without a lateral setoid subapically.Prefemur micropapillate laterally.Caudomedial outgrowth of femur relatively narrow at base (Fig. 2H). Caudomedial outgrowth of tibia with a micropapillate tubercle at base (Fig. 2H). Tarsus quite broadly rounded apically (Fig. 2G, H).

REMARKS This new abundant material, which can be considered as near-topotypic, allows a better understanding of the pronounced morphological variation of this remarkable species.Given the vast distance between Nanjing and South Korea, the recent record of H. emarginata in S Korea by Mikhaljova & Lim (2000) requires verification.