Review of the millipede genus Glyphiulus Gervais, 1847, with descriptions of new species from Southeast Asia (Diplopoda, Spirostreptida, Cambalopsidae). Part 1: the granulatus-group


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Review of the millipede genus Glyphiulus Gervais, 1847, with descriptions of new species from Southeast Asia (Diplopoda, Spirostreptida, Cambalopsidae). Part 1: the granulatus-group

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Review of the millipede genus Glyphiulus Gervais, 1847, with descriptions of new species from Southeast Asia (Diplopoda, Spirostreptida, Cambalopsidae). Part 1: the granulatus-group
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Zoosystema
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Golovatch, Sergei I.
Geoffroy, Jean-Jacques
Mauries, Jean-Paul
Van Den Spiegel, Didier
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Diplopoda ( local )
Cambalopsidae ( local )
Glyphiulus ( local )
Cave ( local )
Southeast Asia ( local )
China ( local )
Laos ( local )
New Species ( local )
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serial ( sobekcm )

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Th e large, widespread, Southeast Asian genus Glyphiulus Gervais, 1847 is reviewed, with special reference to the granulatus-group, established for the species whose primary characteristics are that the males have sternum 1 devoid of median structures, but supplied with two widely separated prongs, at the base of which are often 1- or 2-segmented vestigial legs, along with the carinotaxy patterns of the collum and metaterga which are highly characteristic and often stable. This group currently encompasses 20 species, all keyed here, including the redescribed pantropical species G. granulatus (Gervais, 1847), as well as G. subgranulatus n. sp., G. paragranulatus n. sp., G. semigranulatus n. sp., G. beroni n. sp., G. deharvengi n. sp., G. pergranulatus n. sp., G. latellai n. sp. and G. basalis n. sp., all from caves in southern China, and G. bedosae n. sp. from a cave in Laos. Th e following two synonymies are established: G. granulatus (Gervais, 1847) = G. vulgatus Zhang & Li, 1982, and G. septentrionalis Murakami, 1975 = G. multicarinus Zhang & Li, 1982.
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Zoosystema, Vol. 29, no. 1 (2007).

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7 ZOOSYSTEMA • 2007 • 29 (1) © Publications Scienti Þ ques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com Review of the millipede genus Glyphiulus Gervais, 1847, with descriptions of new species from Southeast Asia (Diplopoda, Spirostreptida, Cambalopsidae). Part 1: the granulatus -groupSergei I. GOLOVATCHInstitute for Problems of Ecology and Evolution, Russian Academy of Sciences,Leninsky pr. 33, Moscow 119071 (Russia) sgol@orc.ruJean-Jacques GEOFFROYMuséum national d’Histoire naturelle, Département Écologie et Gestion de la Biodiversité, USM 306 Réseaux trophiques du sol, 4 avenue du Petit Château, F-91800 Brunoy (France) geoffroy@mnhn.frJean-Paul MAURIÈSMuséum national d’Histoire naturelle, Département Systématique et Évolution, USM 602, case postale 53, 57 rue Cuvier, F-75231 Paris cedex 05 (France) collmill@mnhn.frDidier VAN DEN SPIEGELMusée royal de l’Afrique centrale, Leuvensesteenweg 13, B-3080 Tervuren (Belgium) didier.van.den.spiegel@africamuseum.beKEY WORDS Diplopoda, Cambalopsidae, Glyphiulus , cave, Southeast Asia, China, Laos, new species.Golovatch S. I., Geoffroy J.-J., Mauriès J.-P. & Van den Spiegel D. 2007. — Review of the millipede genus Glyphiulus Gervais, 1847, with descriptions of new species from Southeast Asia (Diplopoda, Spirostreptida, Cambalopsidae). Part 1: the granulatus -group. Zoosystema 29 (1) : 7-49.ABSTRACT e large, widespread, Southeast Asian genus Glyphiulus Gervais, 1847 is reviewed, with special reference to the granulatusgroup, established for the species whose primary characteristics are that the males have sternum 1 devoid of median structures, but supplied with two widely separated prongs, at the base of which are often 1or 2-segmented vestigial legs, along with the carinotaxy patterns of the collum and metaterga which are highly characteristic and often stable. is group currently encompasses 20 species, all keyed here, including the redescribed pantropical species G. granulatus (Gervais, 1847), as well as G. subgranulatus n. sp., G. paragranulatus n. sp., G. semigranulatus n. sp., G. beroni n. sp., G. deharvengi n. sp., G. pergranulatus n. sp., G. latellai n. sp. and G. basalis n. sp., all from caves in southern China, and G. bedosae n. sp. from a cave in Laos. e following two synonymies are established: G. granulatus (Gervais, 1847) = G. vulgatus Zhang & Li, 1982, and G. septentrionalis Murakami, 1975 = G. multicarinus Zhang & Li, 1982.

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8ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. INTRODUCTIONGlyphiulus Gervais, 1847 was “ rst proposed as a subgenus of the then all-embracing genus Iulus Linnaeus, 1758 (recte: Julus ) for a peculiar, strongly ornamented juliform, J. granulatus Gervais, 1847 (Fig. 1), taken on two of the Mascarene islands in the Indian Ocean (Gervais 1847). A similarly strongly-crested juliform was described soon afterwards as a new genus and species, Trachyjulus ceylanicus Peters, 1864, from Sri Lanka (Peters 1864). Karsch (1881) proposed two further species of Glyphiulus , but both have since been shown to belong in other genera, within the same superorder Juliformia, but only very remotely related to Glyphiulus (Krabbe 1982).Pocock (1893, 1894), when describing several further strongly-crested species from Myanmar (Burma), Java and Sumatra, synonymized both Glyphiulus and Trachyjulus with Cambala Say, 1821, a genus created long before for the North American C. annulata Say, 1821. Soon afterwards, however, Pocock (1895) resurrected Glyphiulus for G. granulatus , and Trachyjulus for T. ceylanicus (misspelled as T. ceylonicus ). He also described Cambalomorpha formosa Pocock, 1895, from Hong Kong, added two further species from Myanmar to Cambalomorpha , and proposed the genus Cambalopsis Pocock, 1895 for his Cambala calva Pocock, 1893 (incorrectly written as Cambalopsis calvus ). Since then, Glyphiulus has become one of the most speciose and characteristic genera in the millipede fauna of Southeast Asia, with about 20 species recognized to date, while the genus itself has as many as seven synonyms (Jeekel 2004). ere is no general agreement concerning the scope of the genus Glyphiulus or of the family Cambalopsidae Cook, 1895, to which it belongs. Cambalopsidae are currently divided into three subfamilies (Mauriès 1983, 1992; Jeekel 1985, 2004). Cambalopsinae Cook, 1895 comprise the single but speciose genus Trachyjulus (28 species or subspecies ranging mainly from Sri Lanka, India and Nepal in the West, through Vietnam, ailand and Malaysia up to Sulawesi, Indonesia RÉSUMÉRévision des diplopodes du genre Glyphiulus Gervais, 1847 et description de nouvelles espèces dAsie du Sud-Est (Diplopoda, Spirostreptida, Cambalopsidae). Partie 1 : le groupe granulatus . Glyphiulus Gervais, 1847, genre de Cambalopsidae sud-est asiatique et à très large répartition, est révisé. Une attention particulière est portée au groupe granulatus , établi pour les espèces dont les caractéristiques principales sont, dune part, le sternum 1 du mâle dépourvu de structures médianes mais présentant deux prolongements largement séparés, à la base desquels se trouvent des pattes vestigiales comportant le plus souvent un, parfois deux, articles ; et, dautre part, la structure et la disposition des carènes du collum et des métatergites hautement caractéristiques et souvent stables. Ce groupe comporte 20 espèces qui “ gurent toutes dans une clé. Il inclut lespèce pantropicale G. granulatus (Gervais, 1847), qui est redécrite, ainsi que G. subgranulatus n. sp., G. paragranulatus n. sp., G. semigranulatus n. sp., G. beroni n. sp., G. deharvengi n. sp., G. pergranulatus n. sp., G. latellai n. sp. et G. basalis n. sp., provenant de grottes du sud de la Chine, de même que G. bedosae n. sp. provenant dune grotte du Laos. Les deux synonymies suivantes sont formellement établies : G. granulatus (Gervais, 1847) = G. vulgatus Zhang & Li, 1982 et G. septentrionalis Murakami, 1975 = G. multicarinatus Zhang & Li, 1982.MOTS CLÉS Diplopoda, Cambalopsidae, Glyphiulus , grotte, Asie du Sud-Est, Chine, Laos, espèces nouvelles.

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9 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) in the East). Besides Glyphiulus , the Glyphiuli-nae Chamberlin, 1922 include the genera Dolichoglyphius Verhoe , 1938 (one species with two subspecies from the Ryukyus, Japan), Hypocambala Silvestri, 1897 (11 nominate species, scattered from Mauritius, Seychelles and Comoro Islands in the West, through Vietnam, ailand and Indonesia up to the Loyalty Islands, Bismarck Archipelago and New Caledonia in the East), Plusioglyphiulus Silvestri, 1923 (“ ve species in Cambodia, Malaya and Borneo) and Podoglyphiulus Attems, 1909 (10 species or subspecies from Sri Lanka, India, Nepal and Myanmar). Pericambalinae Silvestri, 1909 are represented by three monotypic genera, Bilingulus Zhang & Li, 1981, Chonecambala Mau-riès & Engho , 2000 and Pericambala Silvestri, 1909, and by the oligotypic genus Parabilingulus Zhang & Li, 1981 (two species), all from China or Vietnam. ese subfamilies, as well as the genera concerned, have been based primarily on di erences in the conformation of the mentum of the gnathochilarium and, to a lesser degree, in anterior gonopod structure, tergal ornamentation and a few other characters (Jeekel 1963, 1985; Mauriès 1970, 1977, 1981, 1983; Ho man 1977). Sometimes the family is even allotted the rank of a superfamily, with the constituent subfamilies thus referred to as families (Mauriès & Nguyen Duy-Jacquemin 1997). ere is no general agreement even as regards the position of the Cambalopsidae in the higher classi“ cation of the Juliformia. Ho man (1980) and Jeekel (1985, 2004) treat the Cambalopsidae as belonging to the order Spirostreptida, whereas Mauriès (1977, 1987, 1992) considers the family to be a member of the order Julida.In short, up to now there has been a lot of confu-sion concerning the status, relationships and scope of the Cambalopsidae, as well as of the subfamilies and genera included therein. For practical reasons alone, the classi“ cation adopted here follows the latest catalogue and review by Jeekel (2004), with a few minor but relevant updates at the speci“ c level.A large number of cambalopsids, mostly caverni-colous, originating from various parts of Southeast Asia have become available to us for study from various sources. Because this material contains numerous new species, many of which represent Glyphiulus s.l., a review of the genus became warranted. e heterogeneity of Glyphiulus has already been noted by Attems (1938) and Mauriès (1970, 1977), who outlined several species groups. Because of this, we have decided to focus “ rst on the distinct group of species closely related to G. granulatus . Strangely enough, this widespread, pantropical species, the type of Glyphiulus , has hitherto been scarcely illustrated. e line drawings of Attems (1900), partly republished by Attems (1926) and reproduced here as Figure 6A-D, seem to be the only meaningful illustrations depicting the male characters of G. granulatus . It is therefore not sur-prising that this species has several junior synonyms, including one proposed as new below. Additional line drawings by Brolemann of female characters (from his unpublished IconographieŽ housed in MNHN), only some of which have been published before (Brolemann 1931), are also given here as Figure 6E-H. e treatment of the remaining Glyphiulus s.l. that do not belong to the granulatusgroup is currently in preparation (Golovatch et al. in press and forthcoming papers). is paper continues the general studies of South-east Asian cave diplopod collections carried out recently in part at the MNHN (Geo roy & Golovatch 2004; Stoev 2004; Stoev & Geo roy 2004; Stoev & Engho 2005; Mauriès 2005; Golovatch et al. 2006a, b). ABBREVIATIONSMCSNV Museo Civico di Storia naturale, Verona; MNHN Muséum national dHistoire naturelle, Paris; MRAC Musée royal de lAfrique centrale, Tervuren; NMNHS National Museum of Natural History, So“ a; SEM scanning electron microscopy;ZMUC Zoological Museum, University of Copen-hagen; ZMUM Zoological Museum, Moscow State University.MATERIAL AND METHODS e material serving as the basis for the present contribution derives from the subterranean

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10ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. collections made by Anne Bedos and Louis De-harveng (MNHN) in Laos and southern China, as well as by both Josiane and Bernard Lips (Vil-leurbanne, France), Petar Beron (NMNHS) and Leonardo Latella (MCSNV) and his collaborators in southern China. e bulk of this material has been deposited in MNHN and NMNHS, with a few voucher specimens shared with the collections of MCSNV, ZMUC and ZMUM.SEM micrographs were taken using a JEOL JSM-6480LV scanning electron microscope. Dry SEM material was coated with gold and removed as such from stubs and returned into alcohol, all samples being kept at MNHN. e carinotaxy formulae in the descriptions below are mainly those proposed by Ho man (1977), but with modi“ cations. e carino-taxy formula of the collum designates the main, uninterrupted crests by Roman numerals, the interrupted ones by Arabic numerals (frontally interrupted marked aŽ, caudally interrupted marked cŽ), the paramedian and median crests are designated by either upper case PŽ or MŽ or by lower case pŽ or mŽ, respectively. is allows not only the number but also the location and shape of the respective crest to be indicated ( cf. Ho man 1977). us, a typical arrangement (from laterad to mesad) and form of the crests on the collum in the species group concerned is as follows: I-VI+7a+pc+ma+pc+7a+VI-I. So it di ers from the simpli“ ed VI-VI and 2-m-2 formula that Ho man (1977) provided for G. granulatus . e carinotaxy pattern of metaterga “ rst shows the number of crests below the ozoporiferous one (designated as IŽ), followed by that be-tween the IŽ. A typical carinotaxy formula of metaterga in the species group concerned is as follows: 3+I+3+I+3 or 2+I+3+I+2. As the crests are usually divided transversely into two halves, while the median crest is often doubled anteriorly, the formula becomes 3(2)+I+4+I+3(2) and 3(2)+i+3+i+3(2), each part corresponding to the number of transverse rows of tubercles (Ho -man 1977). e body segment counts follow Engho et al. (1993), with minor modi“ cations. SYSTEMATICS Order SPIROSTREPTIDA Brandt, 1833 Suborder CAMBALIDEA Bollman, 1893 Family CAMBALOPSIDAE Cook, 1895Cambalopsidae Cook, 1895: 6. Trachyiulidae Silvestri, 1896 (recte: Trachyjulidae): 52. Glyphiulinae Chamberlin, 1922: 2. Agastrophinae Verhoe , 1924: 74. Dolichoglyphiulinae Verhoe , 1938: 88.Genus Glyphiulus Gervais, 1847Glyphiulus Gervais, 1847: 170. Glyphijulus Karsch, 1881: 14. Type species: Iulus (recte: Julus ) granulatus Gervais, 1847; a junior objective synonym, synonymized by Jeekel (1971).Cambalomorpha Pocock, 1895: 363. Type species: Cambalomorpha formosa Pocock, 1895; synonymized by Mauriès (1970).Keratoglyphiulus Attems, 1909: 63. Type species: Iulus (recte: Julus ) granulatus Gervais, 1847; a junior objective synonym, synonymized by Verhoe (1924). Formosoglyphius Verhoe , 1936: 56. Type species: Formosoglyphius tuberculatus Verhoe , 1936; synonymized by Mauriès (1970).Koinoglyphius Carl, 1941: 287. Type species: Glyphiulus su-perbus Silvestri, 1923; synonymized by Jeekel (2004).Octoglyphus Loksa, 1960: 142. Type species: Octoglyphus pulcher Loksa, 1960; synonymized by Murakami (1975).Trogloglyphus Loksa, 1960: 139. Type species: Trogloglyphus balazsi Loksa, 1960; synonymized by Mauriès (1970). TYPE SPECIES. „ Iulus (recte: Julus ) granulatus Gervais, 1847.SPECIES INCLUDED. „ Below is a checklist of the species currently attributed to Glyphiulus , arranged in alphabetical order (after Jeekel 2004). In contrast, Mauriès & Nguyen Duy-Jacquemin (1997) referred to only 16 or 17 species as belonging to this genus, depending on synonymy. 1. G. adeloglyphus Zhang & Li, 1982, described from

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11 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) Xingping, Guangxi Province, China; still only known from the original description (Zhang & Li 1982).2. G. anophthalmus (Loksa, 1960), described as Trogloglyphus anophthalmus Loksa, 1960 from Nyu-Jie Cave near Pulung (= Fulong, near the frontier with Vietnam), Guangxi Province, China; still only known from the original description (Loksa 1960).3. G. balazsi (Loksa, 1960), described as Trogloglyphus balazsi Loksa, 1960 from Lodjen (= Luodian, or Longping), southern Guizhou Province, China; still only known from the original description (Loksa 1960).4. G. capucinus Attems, 1938, described from Mt. Bana, Danang Province, central Vietnam (Engho et al. 2004); still only known from the original description (Attems 1938). 5. G. formosus (Pocock, 1895), described as Cambalomorpha formosa Pocock, 1895 from Hong Kong, China; known from the original description (Pocock 1895) and a partial redescription (Mauriès 1970).6. G. granulatus (Gervais, 1847), described as Iulus (recte: Julus ) ( Glyphiulus ) granulatus Gervais, 1847 from Mauritius and Réunion islands, Indian Ocean; currently known as a pantropical trampŽ species from Hawaii, Hong Kong, Marquesas, Society Islands, Cook Islands, Samoa, Loyalty Islands and New Caledonia in the Paci“ c Ocean, from the Mascarenes, Seychelles and Comoro Islands in the Indian Ocean, and from St. Helena in the Atlantic Ocean (Jeekel 2004). Cambala nodulosa Butler, 1876, from Rodrigues Island, Mascarene Islands, Formosoglyphius tuberculatus Verhoe , 1936, from Taiwan, and Glyphiulus vulgatus Zhang & Li, 1982, from Longzhou, Guangxi Province, China, are junior subjective synonyms of this species (see below).7. G. javanicus Carl, 1911, described from Passaroean, Java, Indonesia; still only known from the original description (Carl 1911).8. G. lipsorum Mauriès & Nguyen Duy-Jacquemin, 1997, described and only known from a cave not in Hubei Province, China, as reported originally (Mauriès & Nguyen Duy-Jacquemin 1997), but in Guangxi Province (Fig. 30).9. G. mediator Attems, 1938, described from Mt. Bana, Danang Province, central Vietnam (Engho et al. 2004); still only known from the original description (Attems 1938).10. G. melanoporus Mauriès & Nguyen Duy-Jacquemin, 1997, described and only known from a cave near Guilin, Guangxi Province, China (Mauriès & Nguyen Duy-Jacquemin 1997). 11. G. pulcher (Loksa, 1960), described as Octoglyphus pulcher Loksa, 1960 from Nyu-Jie Cave near Pulung (= Fulong, near the frontier with Vietnam), Guangxi Province, China; still only known from the original description (Loksa 1960). 12. G. quadrohamatus Chen & Meng, 1991, described from several caves in the Zhenning Bouyzu Miaozu Autonomous County, Guizhou Province, China (Chen & Meng 1991). 13. G. rayrouchi Mauriès & Nguyen Duy-Jacquemin, 1997, described and only known from Hei Yan Cave near Maguan, Guizhou Province, China (Mauriès & Nguyen Duy-Jacquemin 1997). 14. G. recticullus Zhang & Li, 1982, described and still only known from Qinyuan, Zhejiang Province, China (Zhang & Li 1982). 15. G. septentrionalis Murakami, 1975, described from Okinawa Island, Ryukyus, Japan (Murakami 1975), but apparently introduced there from continental China, given that G. multicarinus Zhang & Li, 1982, described from near Guilin, Guangxi Province, China (Zhang & Li 1982), is its junior synonym (see below).16. G. siamensis Mauriès, 1983, described and still only known from Doi Sutep, northern ailand (Mauriès 1983). Engho (2005) records further material from the type locality. 17. G. superbus Silvestri, 1923, described from Dalat, Lamdong Province, central Vietnam (Attems 1938), later recorded from Kampot, Cambodia (Attems 1953), although the latter record needs con“ rmation (Engho et al. 2004). e carinotaxy formula as given in the original description is somewhat ambiguous (Silvestri 1923) and di cult to interpret (see key below). 18. G. vietnamicus Mauriès, 1977, described and only known from a cave in Ke Bang karst, Quang-Binh Province, Vietnam (Mauriès 1977; Engho et al. 2004).19. G. zorzini Mauriès & Nguyen Duy-Jacquemin, 1997, described and still only known from Anjia Yan Cave, Shuichang County, Guizhou Province, China (Mauriès & Nguyen Duy-Jacquemin 1997). e granulatus -group REMARKSSpecies of the granulatusgroup are distinguished by the following characters:1. Male legs 1 are usually very strongly reduced, represented solely or mainly by a sternum lacking any median structures but bearing strongly separated, evidently curved prongs with a leg vestige on each side at base (usually represented by just a few setae, more seldom also one or two rudimentary segments). Very seldom, a nearly complete telopodite persists, but the sternum still supports a pair of widely separated and

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12ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. curved prongs. [Male legs 1 with central structures on the sternum in other groups].2. Other male legs normal or nearly so, not en-larged. [Male legs 2 sometimes incrassate in other groups].3. A typical carinotaxy pattern of the collum is I-VI+7a+pc+ma+pc+7a+VI-I. Quite often the pattern is di erent, either due to crest reduction (especially anteriorly) or hypertrophy, but a median crest is always traceable, at least near the caudal margin. [Usually typical carinotaxy formulae of the collum are di erent in other groups].4. A typical carinotaxy pattern of the metaterga is 3(2)+I+3+I+3(2). e crests usually being divided transversely into two halves, while the median crest is often doubled anteriorly, the formula becomes 3(2)+I+4+I+3(2) and 3(2)+i+3+i+3(2). If the pattern is di erent, it is usually readily derived from the typical one (or vice versa ), i.e. the lateral crests are reduced to two or the crests are undivided, or some of the crests are divided into three, rather than two, transverse rows of tubercles (see also below), etc. A median crest, even when strongly reduced, is always present as well. [Usually typical carinotaxy formulae of the metaterga are di erent in other groups]. 5. e anterior gonopods are reduced to a plate-like coxosternum with moveable, lateral, 1segmented telopodites. [Shared with some other species groups].6. e posterior gonopods are highly compressed, showing a sub” agelliform (rarely subspatuliform), often plumose, distal process. [Shared with some other species groups]. 7. e pleural ” aps behind the gonopod opening on male segment 7 usually do not form an apparent transverse ventral ridge. [Shared with some other species groups]. e group thus roughly corresponds to the concept of Glyphiulus in the sense of Verhoe (1936) and Loksa (1960). At present it contains the follow-ing species: G. adeloglyphus , G. anophthalmus , G. balazsi , G. capucinus , G. granulatus , G. lipsorum , G. melanoporus , G. quadrohamatus , G. rayrouchi , G. septentrionalis , G. superbus , as well as all nine new species described below. e remaining known cambalopsid species are considered as belonging to other groups and will be treated elsewhere. Below a description is given of the available ma-terial, followed by a key to all of the constituent species of the granulatus -group, as well as a brief analysis of their variation and distribution. Glyphiulus granulatus (Gervais, 1847) (Figs 1-6)Iulus granulatus Gervais, 1847: 170, pl. 44, “ g. 10. Glyphiulus granulatus … Karsch 1881: 14. Cambala nodulosa Butler, 1876: 444, synonymized by Jeekel (2004: 53).Formosoglyphius tuberculatus Verhoe , 1936: 57, “ gs 7-9, synonymized by Korsós (2004: 18, 19). Glyphiulus vulgatus Zhang & Li, 1982: 85-87, “ gs 1-6, n. syn.TYPE MATERIAL. „ Réunion (= Île Bourbon). La Bonite , 1838, leg. Eydoux, lectotype (incomplete, lacking caudal body end), 5 paralectotypes (all fragmented), all here designated (MNHN GA 013); material in rather poor condition (most legs missing), apparently dried before being placed in alcohol.Mauriès (1983) mentioned that only part of the type material of this species is still preserved in the MNHN, with the series from Île de France (= Mauritius) presumably being lost. e lectotype selection made here is necessary to “ x the exact type locality (Réunion Island). TYPE LOCALITY. „ Réunion. OTHER MATERIAL EXAMINED. „ Réunion. 10.II.1972, leg. J. Travé, strict topotype juv. (MNHN GA 022).Mauritius. Near Mahébourg, sugar cane debris, 6.II.1972, leg. J. Travé, 1 (det. J.-P. Mauriès, MNHN GA 022).Comoros. Mayotte Island, Mt. Combani, 12°48S, 045°08E, 470 m, Winkler extraction of forest litter, 22-24.II.1999, leg. Jocqué & De Smet, 3 , 1 , 1 juv. (ZMUM, ex coll. MRAC No. 017.940), several and (SEM).China. Hong Kong, leg. M. Crosland, det. J.-P. Mauriès, 2 (MNHN GA 013).Fiji. Viti Levu Island, Suva, park, in litter, 5.VI.1980, leg., det. & don. S. Golovatch, 1 (MNHN GA 013). Seychelles . Mahé Island, 1892, leg. C. Alluaud, det. H. W. Brölemann, 1 complete , 1 incomplete (MNHN GA 013).New Caledonia. Dumbea, leg. Mme Pruvot, det. H. W. Brolemann, 4 , 2 (MNHN GA 013). Loyalty Islands. Lifou, leg. Mme Pruvot, det. H. Brolemann, 1 incomplete (MNHN GA 013). DIAGNOSIS. „ Di ers from congeners primarily by the peculiar anterior gonopods, each showing an unusually high median outgrowth of the coxosternum, combined with the typical patterns of carinotaxy.

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13 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) FIG. 1. — Habitus of Glyphiulus granulatus (Gervais, 1847), from the Comoro Islands (del. N. E. Van Noppen, MRAC).DESCRIPTIONLength of adults of both sexes 10-18 mm, midbody segments round in cross-section (Fig. 3B), their width (horizontal diameter) and height (vertical diameter) similar, 0.6-0.9 mm. Coloration yellow-brown; non-faded specimens variegated, with a dark brown vertex, blackish ocellaria, mainly brown crests on collum and brownish lateral longitudinal stripes beginning from dark brown ozoporiferous tubercles. Sometimes a thin axial line traceable due to darker median crests. Adult body with 30-53 podous segments + 4-2 apodous ones + telson (formula 30-38p+4a+T in smaller specimens, up to 44-53p+2a+T in larger ones; formulae generally following Engho et al. 1993). Clypeus with 3-6 teeth anteromedially. Eye patches transversely ribbon-shaped (smaller specimens) to ovoid (larger individuals), each com-posed of 11-18 rather ” at ocelli in 5 or 6 irregular longitudinal rows (Fig. 4A). Antennae short and clavate (Figs 2B; 3A; 4A), antennomeres 6 and 7 with a small distodorsal “ eld or corolla of ba-cilliform sensilla (Fig. 4B, C). Gnathochilarium usually, but not always (n = 10:1), with a separate promentum (Fig. 3A).Head width = segment 2 < collum = midbody segments (close to 18th to 20th) > segment 3 = 6 > 4 = 5 < 7 < 8 = 10; body abruptly tapering toward telson on a few posteriormost segments. Postcollar constriction very evident (Fig. 2C).Collum with 7+7 longitudinal crests starting from fore edge, but both median crests inter-rupted in about caudal 1/3, being replaced there by similar 1+1+1 crests (Fig. 2A-C) (formula I-VI+7a+pc+ma+pc+7a+VI-I). Subsequent metaterga similarly strongly crested (Fig. 2B-E), especially so from segment 5 on-wards, whence enlarged pore-bearing tubercles commence, less evident on legless segments in front of telson due to loss of ozopores (Fig. 2E). Ozoporiferous tubercles round, wider than high; midbody metatergal crests divided into two about midway, each half neither especially high nor sharp (Fig. 2B-F). Carinotaxy formulae 3+I+4+I+3 and 3+i+3+i+3, the former standing for front row of crests, the latter for caudal one, both fairly independent (Figs 2D-F; 3B).Tegument delicately alveolate-areolate, dull throughout. Fine longitudinal striations in front of stricture between proand metazona, remaining sur-face of prozona very delicately shagreened. Metater-gal setae absent. Segment 2 with long pleural ” aps (Fig. 2B). Limbus extremely “ nely and more or less regularly denticulate. Epiproct simple, devoid

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14ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. A B C D E F FIG. 2. — Glyphiulus granulatus (Gervais, 1847), specimens from the Comoro Islands: A , collum, dorsal view; B , anterior part of body, lateral view; C , same, dorsal view; D , middle part of body, lateral view; E , caudal part of body, lateral view; F , same, dorsal view. Scale bars: A, C, E, F, 0.2 mm; B, 0.5 mm; D, 0.1 mm.of tuberculation, like a transverse rounded ridge in caudal part (Figs 2E, F; 3C, D). Paraprocts rather regularly convex, each with a row of several setae at medial margin and 2+2 setae more laterally (Fig. 3C, D). Hypoproct rather evidently but broadly emarginate caudally to receive ventral edges

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15 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) A B C D E F FIG. 3. — Glyphiulus granulatus (Gervais, 1847), specimens from the Comoro Islands: A , head, ventral view; B , midbody segment section, caudal view; C , caudal part of body, ventral view; D , same, caudal view; E , segments 6-8 of , ventral view; F , segment 7 of , frontolateral view. Scale bars: A, B, 0.2 mm; C-F, 0.1 mm.

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16ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. AB C D EF FIG. 4. — Glyphiulus granulatus (Gervais, 1847), specimens from the Comoro Islands: A , cephalic capsule, sublateral view; B , antenna, apical view; C , bacilliform sensilla on antennomere 6, lateral view; D , male legs 1, caudal view; E , male legs 2, caudal view; F , male legs 3, frontal view. Scale bars: A, D-F, 0.1 mm; B, 0.05 mm; C, without scale.of paraprocts, with 1+1 strongly separated setae near caudal margin (Fig. 3C).Ventral ” aps behind gonopod opening on male segment 7 barely distinguishable as low swell-ings, not forming a marked transverse ridge (Fig. 3E, F).Legs short, on midbody segments about 3/4 length of segment height (Figs 2B, D, E; 3B). Claw at base with a strong accessory spine almost half as long as claw itself (Fig. 5D). Tarsi characteristically fringed

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17 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) AB CD FIG. 5. — Glyphiulus granulatus (Gervais, 1847), specimens from the Comoro Islands: A , anterior gonopods, frontal view; B , same, caudal view; C , posterior gonopods, frontal view; D , claw of midbody leg, lateral view. Scale bars: A-C, 0.05 mm; D, 0.01 mm.terminally, some of terminal setae with scattered denticles (Fig. 5D). Male legs 1 highly characteristic (Figs 4D; 6A) in being very strongly reduced, represented only by a sternum devoid of any median or parame-dian structures but carrying 1+1 strongly sepa-rated prongs, both evidently curved posteriad and bearing several strong setae and a minute tubercle (vestige of legs) at base on caudal face. Male legs 2 very slightly hypertrophied, only claw and, anteriorly, coxa somewhat reduced; penes broad, oblong-subtrapeziform, each with 2 or 3 strong setae distolaterally (Figs 4E; 6B). Male legs 3 modi“ ed in having coxa especially slender and elongate (Fig. 4F).Anterior gonopods (Figs 5A, B; 6C) with a typical shield-like coxosternum which is modestly setose on caudal face and provided with a pair of con-spicuous, high, terminally coiled, mesal processes (visible also in situ , Fig. 3E). Telopodite small but movable, 1-segmented, lateral in position, with 2 or 3 strong apical setae and a “ eld of small setae at base, slightly longer than adjacent lateral corner of coxosternum. Posterior gonopods (Figs 5C; 6D) very compact, coxite medio-apically with a plumose distal process and a hyaline lobe, telopodite setose both laterally and medially, lower than both coxal process and lobe.Vulvae very simple, bare, modestly emarginate medially (Fig. 6G, H). Female legs 1 and 2 as in Figure 6E, F, female coxa 3 as slender as in male (Fig. 6G). Early juvenile stadia recognised by normal, large ozoporiferous tubercles on segments 5 and 6, these tubercles being considerably reduced in size on subsequent segments.

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18ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. A B C D EF H G FIG. 6. — Glyphiulus granulatus (Gervais, 1847), ( A D ), ( E H ): A , legs 1; B , legs 2; C , anterior gonopods, caudal view; D , posterior gonopods, caudal view; E , legs 1; F , legs 2, caudal view; G , vulvae with base of legs 3, caudal view; H , left vulva, frontal view. Drawn not to scale. A-D, after Attems (1900); E, G, after Brolemann’s “Iconographie” (kept in MNHN); F, H, after Brolemann (1931).

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19 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) AB CD E F FIG. 7. — Glyphiulus subgranulatus n. sp., paratype: A , head, ventral view; B , collum, dorsal view; C , middle part of body, dorsal view; D , same, lateral view; E , caudal part of body, lateral view; F , same, dorsal view. Scale bars: A, 0.1 mm; B, 0.2 mm, C-F, 0.5 mm.REMARKS e above is the “ rst formal record of G. granulatus in Fiji.Based on the original description and illustra-tions of G. vulgatus alone (Zhang & Li 1982), it is clear that this taxon does not di er in any way from G. granulatus , thus justifying the above new synonymy. e far-inland record of G. granulatus (= G. vulgatus ) in Guangxi Province, southern China suggests a continental Chinese origin of this species, which has since become established on numer-ous tropical islands all over the globe. e port of Hong Kong seems to have been a plausible outlet for the recent expansion of G. granulatus through human agency. e fact that, of all the numerous congeners, only this species that has attained such

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20ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. A B C D FIG. 8. — Glyphiulus subgranulatus n. sp., and paratypes: A , anterior part of body, lateral view; B , caudal part of body, ventral view; C , midbody segment section (a little damaged), caudal view; D , male legs 1, caudal view. Scale bars: A, C, 0.5 mm; B, 0.2 mm; D, 0.1 mm.a vast distribution seems to be rooted in its biology and ecology, the most relevant traits being its fairly small size (on average the smallest) and, ap-parently, a relatively rapid development, suggested indirectly by the observations of Engho (1993) and Mauriès & Nguyen Duy-Jacquemin (1997). e ontogeny of certain cambalopsids is unique among Diplopoda in containing a haplopodous early juvenile stage (third?). At this stage, all body segments after the sixth are only equipped with a single pair of legs! In G. granulatus , the total number of such segments is only up to 13, as opposed to 18 in G. subgranulatus n. sp. and G. semigranulatus n. sp. (see below), 19 in Trachyjulus tjampeanus (Attems, 1903), from Java, Indonesia (NB: in the abundant material of cambalopsids taken from numerous caves from all over Java and accumu-lated at MNHN, we have only encountered this species!), and 28 in Glyphiulus zorzini , a southern Chinese cavernicole. In addition, in the Hawaiis, G. granulatus has been recorded as a myrmecophile (Crosland 1994).Similarly, based solely on the original descrip-tions of G. septentrionalis and G. multicarinus ( cf. Murakami 1975; Zhang & Li 1982), it is clear that these species cannot be separated, agreeing perfectly with each other in every character. Hence the new synonymy advanced: G. septentrionalis Murakami, 1975 = G. multicarinus Zhang & Li, 1982. e oc-currence of this species on Okinawa, Ryukyus, both in caves and epigeically (Murakami 1975), seems only to represent a recent introduction through hu-man agency, with southern China de“ nitely having served as the source area. Much like G. granulatus ,

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21 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) ABC D E F G FIG. 9. — Glyphiulus subgranulatus n. sp., paratype: A , antenna; B , gnathochilarium, ventral view; C , legs 1, caudal view; D , legs 2, caudal view; E , anterior gonopods, caudal view; F , posterior gonopods, frontal view; G , same, caudal view. Scale bar: A, B, 0.3 mm; C-G, 0.2 mm.G. septentrionalis is a relatively small species (adults 17-30 mm long), but the number of segments is usually considerably higher (43-66p+4-1a+T), implying a longer development, which apparently reduces its vagility. Glyphiulus subgranulatus n. sp. (Figs 7-9)TYPE MATERIAL. „ China. Yunnan Prov., Mengzi County, cave near footpath to plateau, 5.I.1989, leg. P. Beron, holotype (NMNHS); paratypes 7 , 14 , 11 juv. (NMNHS); 2 , 2 (MNHN GA 035); 2 , 2 (ZMUM); 1 , 1 (ZMUC); 1 , 1 (MCSNV); 1 , 1 (SEM). „ Yunnan Prov., Mengzi County, Pothole No. 2 (Ma Fa Tiao Dong), 6.I.1989, leg. P. Beron, paratype 1 (NMNHS).ETYMOLOGY. „ To emphasize the obvious similarity to G. granulatus .DIAGNOSIS. „ Di ers from the most similar congeners by the crests on the collum being underdeveloped (especially so anteriorly), the body segments somewhat compressed laterally, the anterior gonopods with unusually slender telopodites, etc.DESCRIPTIONLength of adults of both sexes 19-28 mm, width 1.0-1.3 mm; males usually a little shorter and more slender than females. Coloration much as in G. granulatus , variegate, yellow-brown, with a dark brown vertex, blackish ocellaria, mainly brown crests on collum and grey-brownish lateral longitudinal stripes running down from dark brown ozoporiferous tubercles; dorsum generally light, but a thin axial line mostly present due to darker median crests.Adults with 38-52p+5-2a+T in males, 39-63p+5-1a+T in females; largest juvenile with 31p+3a+T; two smallest juveniles with 17 or 18 haplopodous

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22ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. A B C D E F FIG. 10. — Glyphiulus paragranulatus n. sp., and paratypes: A , anterior part of body, lateral view; B , same, dorsal view; C , middle part of body, dorsal view; D , same, lateral view; E , caudal part of body, dorsal view; F , same, caudal view. Scale bars: A, D, 0.5 mm; B, C, E, F, 0.2 mm.segments after 6th, plus 2a+T. Holotype about 23 mm long, 1.0 mm wide, with 47p+2a+T. Other characters as in G. granulatus , including mentum either divided (n = 1) (Fig. 9B) or un-divided (n = 1) (Fig. 7A), 11-18 ocelli, antennae (Figs 8A; 9A), the carinotaxy formulae (Fig. 7B-F), claw, posterior gonopod structure (Fig. 9F, G), male legs 1 and 2 (Figs 8D; 9C, D), telson (Fig. 8B), etc., but body size much larger (see above); segments compressed laterally (Fig. 8C); crests on collum (7+m+7 at caudal margin) evidently reduced in size, almost obsolete anteromedially (Figs 7B; 8A); legs somewhat longer (about 4/5 as long as body height); anterior gonopods with telopodites more

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23 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) A B C D E F FIG. 11. — Glyphiulus paragranulatus n. sp., and paratypes: A , caudal part of body, ventral view; B , midbody segment section, caudal view; C , midbody leg; D , claw; E , posterior gonopods in situ in gonopod opening, frontal view; F , anterior gonopods, caudal view. Scale bars: A, 0.2 mm; B, 0.5 mm; C, E, F, 0.1 mm; D, 0.02 mm.slender, while the coxosternum has much shorter distomedian outgrowths and acute distolateral corners (Fig. 9E). REMARKSIn the absence of evident troglomorphic traits, this species can only be considered as troglophilic.

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24ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. ABC D E F G H FIG. 12. — Glyphiulus paragranulatus n. sp., paratype: A , antenna; B , gnathochilarium, ventral view; C , legs 1, caudal view; D , legs 2, caudal view; E , leg 3; F , anterior gonopods, caudal view; G , posterior gonopods, frontal view; H , tip of distal process, caudal view. Scale bar: A, B, 0.3 mm; C-G, 0.2 mm.Glyphiulus paragranulatus n. sp. (Figs 10-12)TYPE MATERIAL. „ China. Yunnan Prov., Jianshui County, Yan Dong Cave, 12.I.1989, leg. P. Beron, holotype (NMNHS); paratypes 2 , 1 fragm. juv. (NMNHS); 1 , 1 (MNHN GA 036); 1 (ZMUM); 1 , 1 (SEM).ETYMOLOGY. „ To emphasize the obvious similarity to G. granulatus .DIAGNOSIS. „ Di ers from the most similar congeners by the antennae being more slender, the body seg-ments somewhat compressed laterally, the ocellaria only slightly reduced, the accessory spine at the claw base especially prominent, the anterior gonopod coxosternum with small but evident distomedial outgrowths.DESCRIPTIONLength of adults of both sexes 18-25 mm, width 1.0-1.2 mm. General coloration pallid, pattern much as in G. granulatus , but ocellaria light brown while lateral stripes level to ozoporiferous tubercles narrow.Adults with 39-45p+2-1a+T. Holotype about 25 mm long, 1.2 mm wide, with 44p+2a+T. Other characters as in G. granulatus , including gnathochilarium (divided into proand eumentum, n = 1) (Fig. 12B), the carinotaxy formulae (Figs 10A-D; 11B), telson (Figs 10E, F; 11A), male legs 1-3 (Fig. 12C-E), posterior gonopod structure (Figs 11E; 12G, H), etc., but body size much larger (see above); segments compressed laterally (Fig. 11B); antennae slightly elongate (Figs 10A, B; 12A); claw with a particularly prominent spine at base (Fig. 11D); only some 9-11 ocelli in adults; legs somewhat longer (about as long as body height) (Figs 10D; 11C); anterior gonopod coxosternum with small but curved distomedian outgrowths and acute distolateral corners (Figs 11F; 12F). REMARKSEven though there are some indications of troglomorphism in this species (pallid general coloration, somewhat elongated antennae and legs), its troglophily seems more likely than troglobiosis. e morpho-logical evidence being inconclusive, without further

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25 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) A B CD EF FIG. 13. — Glyphiulus semigranulatus n. sp., and paratypes: A , anterior part of body, lateral view; B , same, dorsal view; C , midbody segment section, caudal view; D , middle part of body, dorsal view; E , caudal part of body, dorsal view; F , telson, caudal view. Scale bars: A, D, 0.5 mm; B, C, E, 0.2 mm; F, 0.1 mm.observations this problem can never be solved. On the other hand, Bollmania beroni Stoev & Engho , 2005 (Diplopoda, Callipodida, Caspiopetalidae Lohmander, 1931), a presumed troglobitic millipede, occurs in the same cave (Stoev & Engho 2005). Glyphiulus semigranulatus n. sp. (Figs 13; 14)TYPE MATERIAL. „ China. Yunnan Prov., Meele (= Mile) County, Bai Long Dong Cave (= White Dragon Cave),

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26ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. A B CD EF G FIG. 14. — Glyphiulus semigranulatus n. sp., paratype: A , antenna; B , gnathochilarium, ventral view; C , legs 1, caudal view; D , leg 3; E , anterior gonopods, caudal view; F , posterior gonopods, caudal view; G , same, frontal view. Scale bar: 0.2 mm.2.I.1989, leg. P. Beron, holotype (NMNHS); paratypes 1 , 1 (NMNHS); 1 (SEM); 1 (MNHN GA 037). „ Yunnan Prov., Jianshui County, Yan Zi Dong Cave, 10.I.1989, leg. P. Beron, paratypes 5 , 3 , 4 juv. (NMNHS); 4 , 2 (MNHN GA 037); 4 , 2 (ZMUM); 2 , 2 (ZMUC); 1 , 1 (MCSNV); 1 (SEM).ETYMOLOGY. „ To emphasize the obvious similarity to G. granulatus .DIAGNOSIS. „ Di ers by a position somewhat intermedi-ate between the most similar congeners G. granulatus and G. paragranulatus n. sp., i.e. coloration, carinotaxy, antennae, claws, etc., as in G. granulatus , but size, segment shape, leg length and anterior gonopods more like in G. paragranulatus n. sp.DESCRIPTIONLength of adults of both sexes 12-24 mm, width 0.8-1.2 mm. Adults of both sexes with 31-64p+5-1a+T; largest juvenile with 28p+5a+T, largest juvenile with 27p+5a+T; two smallest juveniles with 15 or 18 haplopodous segments after 6th, plus 2a+T or

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27 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) AB C D E F FIG. 15. — Glyphiulus beroni n. sp., paratype: A , anterior part of body, lateral view; B , same, dorsofrontal view; C , middle part of body, dorsal view; D , same, lateral view; E , caudal part of body, dorsal view; F , same, ventral view. Scale bars: A-E, 0.5 mm; F, 0.2 mm.1a+T, respectively. Length of holotype about 22 mm, width 1.0 mm, with 45p+3a+T. Coloration, carinotaxy (Fig. 13A-E), antennae (Figs 13A; 14A), gnathochilarium, claws, etc. (Fig. 14B-D, F, G) basically as in G. granula-tus , but body size (usually much larger), segment shape (slightly compressed laterally) (Fig. 13C), telson (Fig. 13E, F) with peculiar ” aps on parap-rocts near hypoproct (Fig. 13F), legs (somewhat longer) and anterior gonopods (Fig. 14E) more like those of G. paragranulatus n. sp. Adults with 11-15 ocelli.

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28ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. A B C D EF FIG. 16. — Glyphiulus beroni n. sp., paratype: A , antenna; B , legs 1, caudal view; C , legs 2, caudal view; D , anterior gonopods, caudal view; E , posterior gonopods, caudal view; F , same, frontal view. Scale bar: 0.2 mm.REMARKSIn the absence of any evident troglomorphic traits, this species can only be considered as troglophilic at most. is is supported by G. semigranulatus n. sp. being present in several relatively strongly isolated caves, sometimes in signi“ cant numbers. Glyphiulus beroni n. sp. (Figs 15; 16)TYPE MATERIAL. „ China. Yunnan Prov., Jianshui County, Cave Ba Guo Dong, 11.I.1989, leg. P. Beron, holotype (NMNHS); paratypes 2 , 6 , 3 juv. (NMNHS); 2 , 2 (MNHN GA 038); 1 , 1 (ZMUM); 1 , 1 (ZMUC); 1 (SEM). ETYMOLOGY. „ Honours Petar Beron, the collector.DIAGNOSIS. „ Di ers by the crests just below and above the ozopore-bearing crests being divided into three, not two, transverse rows of tubercles, coupled with an evident mediodorsal tubercle present in front of the caudal edge of the epiproct.DESCRIPTIONLength of adults 11-17 ( ) or 12-24 mm ( ), width 0.8-1.0 ( ) or 0.8-1.3 mm ( ). Coloration nearly entirely light yellowish, only ocellaria brown,

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29 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) AB C D EF FIG. 17. — Glyphiulus bedosae n. sp., paratype: A , anterior part of body, lateral view; B , same, dorsal view; C , middle part of body, lateral view; D , same, dorsal view; E , caudal part of body, lateral view; F , telson, ventral view. Scale bars: A, C, 0.5 mm; B, D, E, 0.2 mm; F, 0.1 mm.as well as sometimes a vague lateral line level to ozoporiferous crests.Adults with 28-40p+5-2a+T ( ) or 28-53p+5-1a+T ( ); largest juvenile with 27p+5a+T, largest juvenile with 27p+4a+T. Length of holotype about 17 mm, width 1.0 mm, with 40p+2a+T. Carinotaxy of collum typical (Fig. 15A, B), that of trunk segments generally typical but, starting from segment 7 until 6 or 7 penultimate segments, crests just below as well as between ozopores divided into three, not two, transverse rows of tubercles (Fig. 15C, D). An evident middorsal tubercle just above caudal edge of epiproct (Fig. 15E).

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30ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. A B C DEF G H FIG. 18. — Glyphiulus bedosae n. sp., paratype: A , antenna; B , gnathochilarium, caudal view; C , legs 1, caudal view; D , legs 2, caudal view; E , leg 3; F , midbody leg; G , anterior gonopods, caudal view; H , posterior gonopod, subcaudal view. Scale bar: A-F, 0.2 mm; G, H, 0.1 mm.Antennae (Figs 15A, B; 16A), legs (Fig. 16B, C), claws, posterior gonopods (Fig. 16E, F), etc., as in G. granulatus , but body size (often much larger), segment shape (slightly compressed laterally) and anterior gonopods (Fig. 16D) more like those of G. semigranulatus n. sp. Adults with 5-13 ocelli. Male legs 1 with tubercular leg vestiges near base of prongs (Fig. 16B). REMARKSIn the absence of any clearly troglomorphic traits, except perhaps for its nearly completely unpigmented body and the somewhat reduced ocellaria, this species can hardly be considered as more than troglophilic. Glyphiulus bedosae n. sp. (Figs 17; 18)TYPE MATERIAL. „ Laos. Luang Prabang Prov., Nong Kiaw (= Muang Ngoy), am Pha Kouang Cave, 29.XII.2000, leg. A. Bedos & L. Deharveng (LAO-099), holotype (MNHN GA 039); paratypes 1 (MNHN GA 039), 1 (SEM). ETYMOLOGY. „ Honours Anne Bedos, one of the collectors.

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31 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) AB CD EF FIG. 19. — Glyphiulus deharvengi n. sp., paratype: A , anterior part of body, lateral view; B , same, dorsal view; C , middle part of body, dorsal view; D , telson, ventral view; E , midbody segment section, caudal view; F , claw of a midbody leg. Scale bars: A, C, D, 0.5 mm; B, E, 1.0 mm; F, 0.1 mm.

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32ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. A B C D E F G H FIG. 20. — Glyphiulus deharvengi n. sp., paratype: A , antenna; B , gnathochilarium, caudal view; C , legs 1, caudal view; D , legs 2, caudal view; E , leg 3; F , midbody leg; G , anterior gonopods, caudal view; H , posterior gonopod, subfrontal view. Scale bar: A-F, 0.6 mm; G, H, 0.3 mm.DIAGNOSIS. „ Di ers from other species by the paraprocts showing a clear median marginal ridge, combined with a plumose spatuliform distal process of the posterior gonopods and a subunciform distomedian corner of the anterior gonopods.DESCRIPTIONLength of adults of both sexes 16-20 mm, width 1.0-1.1 mm.Holotype 16 mm long, 1.0 mm wide, with 40p+3a+T and 11 ocelli; paratype about 20 mm long, 1.1 mm wide, with 54p+2a+T and 14 ocelli. Coloration, antennae (Figs 17A; 18A), gnatho-chilarium (mentum divided, n = 1) (Fig. 18B), carinotaxy (Fig. 17A-E), segment shape, legs (Fig. 18C-F), etc., generally as in G. granulatus , but paraprocts with a low, but evident, median marginal ridge (Fig. 17F).Anterior gonopods like those of G. semigranulatus n. sp. (Fig. 18G), but posterior gonopods with a plumose spatuliform distal process (Fig. 18H), i.e. much as in G. quadrohamatus or G. lipsorum . REMARKSIn the absence of any troglomorphic traits, this species can only be considered troglophilic at most. Glyphiulus deharvengi n. sp. (Figs 19; 20)TYPE MATERIAL. „ China . Hunan Prov., Longshan: Huoyan, Feihu Dong Cave (Flying Tiger Cave), 11.VIII.1995, leg. A. Bedos & L. Deharveng (CHI-022), holotype (MNHN GA 040); paratype 1 (MNHN GA 040). „ Hunan Prov., Longshan, Huoyan, Baiyan Dong Cave (White Rocks Cave), 15.VIII.1995, leg. A. Bedos & L. Deharveng (CHI-032), paratypes 1 , 1 , 1 juv. (MNHN GA 040); 1 (ZMUM). „ Hunan Prov., Longshan: Huoyan,

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33 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) AB CD FIG. 21. — Glyphiulus pergranulatus n. sp., paratype: A , anterior part of body, lateral view; B , same, dorsal view; C , middle part of body, lateral view; D , same, dorsal view. Scale bars: 0.5 mm.Renmi Dong Cave, 19.VIII.1995, leg. A. Bedos & L. Deharveng (CHI-043); paratypes 1 (MNHN GA 040); 1 (SEM). ETYMOLOGY. „ Honours Louis Deharveng, one of the collectors.DIAGNOSIS. „ Di ers by the relatively large size, the nearly completely smooth collum, the complete depigmentation (even of ocellaria), the elongated legs (with claws devoid of a spine at base from leg 4 onwards), the low, but evident, transverse pleural ridge behind the gonopod opening on male segment 7, the bare but ” agelliform distal process of the posterior gonopods, etc.DESCRIPTIONLength of adults of both sexes 40-55 mm, width 2.0-2.3 mm. Coloration usually uniformly dark brown due to heavy sclerotization, tegument being coloured only due to chitin; sometimes ozoporiferous tubercles reddish.Adults with 55-68p+4-1a+T; juvenile with 40p+5a+T, recognised by considerably reduced ozoporiferous tubercles on segments 7-33, plus 5a+T. Length of holotype about 55 mm, width 2.3 mm, with 67p+1a+T.Antennae slightly more elongate than usual (Figs 19A; 20A). Ocellaria unpigmented, barely traceable, with about 10 or 11 ocelli in largest specimens. Gnathochilarium (Fig. 20B) with an undivided mentum (n = 1).Carinotaxy of collum unusual in that all crests are nearly obliterated (Fig. 19A, B), though a typi-cal pattern is traceable. Carinotaxy of subsequent metaterga typical (Fig. 19A-C, E), but crests low, especially dorsal ones on segments 2-6; only two lateral crests below ozoporiferous ones, divided from segment 7 onwards, followed by a dozen striae. A low, but evident, transverse pleural ridge behind gonopod opening on male segment 7. Epiproct slightly elongate apically, more narrowly rounded

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34ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. A B CD FIG. 22. — Glyphiulus pergranulatus n. sp., paratype: A , caudal part of body, ventral view; B , same, lateral view; C , claw of a midbody leg; D , midbody segment section, caudal view. Scale bars: A, 0.2 mm; B, D, 0.5 mm; C, 0.01 mm.than usual; hypoproct more narrowly emarginate at caudal margin (Fig. 19D).Legs elongate (Fig. 20F), claws strongly elon-gate, without a spine near base starting from leg 4 (Fig. 19F).Male legs 1 with vestigial tubercular coxites (Fig. 20C). Penes with 4 or 5 long setae distola-terally (Fig. 20D), male legs 3 elongate, much as in G. granulatus (Fig. 20E). Anterior gonopods with small subunciform protuberances at median corners of coxosternum (Fig. 20G). Posterior gonopods with a ” agelliform but bare distal process (Fig. 20H). REMARKSBased on its evident troglomorphic traits, this species seems to be a troglobite. Glyphiulus pergranulatus n. sp. (Figs 21-23)TYPE MATERIAL. „ China. Guizhou Prov., Guanling County, Huajiang, Da Dong Cave, 4.IV.2004, leg. L. Latella & Ghen Hu, holotype (MNHN GA 041); paratypes 1 (MNHN GA 041); 1 (SEM). „ Guizhou Prov., Guanling County, Huajiang, An Jia Da Dong Cave, 4.IV.2004, leg. L. Latella & R. Zorzin, paratype 1 (MNHN GA 041).ETYMOLOGY. „ To emphasize the close a nities to G. granulatus .DIAGNOSIS. „ Characterized by the somewhat larger ocellaria, the underdeveloped tergal crests (becoming almost obliterated in the median part of the collum), coupled with the medium body size, slightly elongated legs, simple anterior gonopods, etc.

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35 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) AB C D E F G H FIG. 23. — Glyphiulus pergranulatus n. sp., paratype: A , antenna; B , legs 1, caudal view; C , legs 2, caudal view; D , leg 3; E , midbody leg; F , anterior gonopods, caudal view; G , posterior gonopod, subfrontal view; H , same, frontal view. Scale bar: 0.2 mm.DESCRIPTIONLength of holotype and of paratype from An Jia Da Dong about 30 mm, of paratype about 33 mm, width 1.5, 1.5 and 1.7 mm, with 56p+2a+T, 67p+1a+T and 57p+2a+T, respectively. Coloration, antennae (Figs 21A; 23A), segment shape (Fig. 22D), carinotaxy patterns (Fig. 21), male legs 1-3 (Fig. 23B-D), etc., as in G. granulatus , but tergal crests somewhat underdeveloped (Figs 21; 22B, D), almost obliterated in median part of collum (Fig. 21A, B). Ocellaria rounded-triangular, black, each with 23-25 poorly convex ocelli. Claws slightly longer, spine at base slightly shorter (Fig. 22C); legs about 4/5 as long as midbody height (Fig. 23E). Epiproct with a faint but evident middorsal tubercle (Fig. 22B); hypoproct more narrowly emarginate at caudal margin (Fig. 22A).Anterior gonopod coxosternum particularly simple (Fig. 23F). Posterior gonopods with a typical ” agel-liform and plumose distal process (Fig. 23G, H). REMARKSEven though taken from caves, this species seems to represent only a troglophile at most. However, this cave harbours another, apparently troglobitic congener from a di erent species group, to be described elsewhere. Glyphiulus latellai n. sp. (Figs 24-26)TYPE MATERIAL. „ China. Guizhou Prov., Gianxi County, Hong Lin Village, Chang Tu Dong Cave, 13.XI.2003, leg. L. Latella, holotype (MNHN GA

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36ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. AB C D FIG. 24. — Glyphiulus latellai n. sp., paratype: A , anterior part of body, lateral view; B , same, dorsal view; C , middle part of body, dorsal view; D , caudal part of body, lateral view. Scale bars: 0.5 mm.042); paratypes 2 , 1 (MNHN GA 042), 1 (SEM). „ Same locality, 3.IV.2004, leg. L. Latella, paratype 1 ( MCSNV). „ Guizhou Prov., Gianxi County, Hong Lin Village, Xiao Dong Cave, 16.XI.2003, leg. Latella & Cioperelli-Repetto, paratype 1 (ZMUM). „ Guizhou Prov., Gianxi County, Hong Lin Village, Xi Xiang Dong Cave, 14.XI.2001, leg. L. Latella, paratype 1 (MNHN GA 042). „ Guizhou Prov., Gianxi County, Hong Lin Village, Dayan Dong Cave, 22.XI.2003, leg. L. Latella, paratype 1 (ZMUC). „ Guizhou Prov., Gianxi County, Hong Lin Village, Tiao Shuz Dong Cave, 18.XI.2001, leg. L. Latella, D. Avesani & G. Rossi, paratypes 1 , 3 juv. (MNHN GA 042); 1 (SEM). „ Guizhou Prov., Gianxi County, Hong Lin Village, Lu Diao Ai Dong Cave (= Cow Fell Down Cave), XI.2003, leg. L. Latella, paratype 1 (NMNHS). „ Guizhou Prov., Gianxi County, Hong Lin Village, Shu Jia Yan Dong Cave, 17.XI.2001, leg. Latella & Berzacola, paratypes 2 , 1 , 1 juv. (MNHN GA 042). „ Guizhou Prov., Gianxi County, Ishui Luo Dong Cave, 20.XI.2001, leg. L. Latella, paratype 1 (MNHN GA 042).ETYMOLOGY. „ Honours Leonardo Latella, the col-lector.DIAGNOSIS. „ Di ers from congeners by the strongly reduced crests on the collum and metaterga, the completely undivided dorsal crests (the mediodorsal ones also undivided anteriorly), the complete depig-mentation (even of ocellaria), the evidently elongated antennae and legs, the low, but evident, transverse pleural ridge behind the gonopod opening on male segment 7, etc.DESCRIPTIONLength of adults of both sexes 18-45 mm, width 1.3-2.0 mm. Coloration usually uniformly dark brown due to heavy sclerotization, sometimes ozo-poriferous tubercles reddish; more rarely, coloration uniform yellowish. Adults of both sexes with 42-63p+6-1a+T; larg-est juveniles ( ) with 35p+4a+T or 36p+5a+T.

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37 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) AB CD FIG. 25. — Glyphiulus latellai n. sp., paratype: A , telson, ventral view; B , midbody segment section, frontal view; C , bacilliform sensilla on antennomere 6; D , claw of a midbody leg. Scale bars: A, 0.2 mm; B, 0.5 mm; C, 0.05 mm; D, 0.02 mm.Length of holotype about 30 mm, width 1.3 mm, with 50p+3a+T.Antennae much more elongate than usual (Figs 24A; 26A), with especially numerous and dense distodorsal sensilla on antennomeres 5 and 6 (Fig. 25C). Ocellaria, if any, unpigmented, ocelli, when present, 1-11, irregularly spaced, barely trace-able. Gnathochilarium usual, mentum divided (n = 2) (Fig. 26B).Carinotaxy of collum rather typical, though hardly traceable because of crests being nearly obliterated, especially anteromedially (Fig. 24A, B). Carinotaxy of subsequent metaterga deviating slightly from typical formula (Figs 24; 25B) in that none of the dorsal crests is divided transversely or longi-tudinally; all crests somewhat reduced (Fig. 25B), with only 2 or 3 very low lateral ones (formula 2(3)+I+3+I+2(3)). From segment 7 on, crest below IŽ (i.e. below ozoporiferous tubercle) poorly di-vided at about midway; further ventrad, segments with 1 or 2 barely distinguishable crests followed by about a dozen “ ne striae (Fig. 24A, D). A low, but evident, transverse pleural ridge present behind gonopod opening on male segment 7. Epiproct slightly elongate apically, more narrowly rounded than usual; hypoproct more narrowly emarginate at caudal margin (Fig. 25A). Legs elongate (Fig. 25B), about as long as midbody height; claws strongly elongate, with a small spine near base on legs of anterior 1/5-1/4 body (Fig. 26F), thereafter devoid of any basal spines (Fig. 25D). Male legs 1 with rudimentary tubercular

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38ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. A B C DE FGH I J K FIG. 26. — Glyphiulus latellai n. sp., paratype: A , antenna; B , gnathochilarium, caudal view; C , legs 1, caudal view; D , legs 2, caudal view; E , leg 3; F , leg 11; G , anterior gonopods, caudal view; H , left posterior gonopod, caudal view; I , same, frontal view; J , right posterior gonopod, submedial view; K , same, sublateral view. Scale bar: 0.2 mm.coxites (Fig. 26C); male legs 2 and 3 typical (Fig. 26D, E).Anterior gonopods with small, subunciform pro-tuberances at median corners of coxosternum (Fig. 26G). Posterior gonopods with a ” agelliform and plumose distal process (Fig. 26H-K). REMARKSBased on its troglomorphic traits, this species seems to be a troglobite. It coexists in four of the caves

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39 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) AB CD FIG. 27. — Glyphiulus basalis n. sp., paratype: A , anterior part of body, lateral view; B , same, dorsal view; C , middle part of body, lateral view; D , same, dorsal view. Scale bars: A, B, 0.5 mm; C, D, 1 mm.with still another millipede troglobite, Guizhousoma latellai Mauriès, 2005 (Chordeumatida, Guizhou-somatidae) (Mauriès 2005). Glyphiulus basalis n. sp. (Figs 27-29)TYPE MATERIAL. „ China. Sichuan Prov., Xin Long County, Gan Chuan Cave, Nos 414, 417 and 421, 6.VIII.1999, leg. J. & B. Lips, holotype (MNHN GA 043); paratypes 1 , 3 (MNHN GA 043); 1 (ZMUM); 1 (ZMUC); 1 (SEM).ETYMOLOGY. „ To emphasize the presumed basal position of this species within the group.DIAGNOSIS. „ Di ers primarily by the retained telo-podites of male legs 1, coupled with the large size, the crests on the collum and the spines at claw base both strongly reduced, the crests on metaterga reduced but the ozoporiferous tubercles high, the troglomorphic traits (complete depigmentation, the lack of ocelli, the evidently elongated antennae and legs), the low, but evident, transverse pleural ridge behind the gonopod opening on male segment 7.DESCRIPTIONLength of , 30-32 mm, width 1.8 mm; of , 23-42 and 1.5-2.2 mm, respectively. Length of holotype about 30 mm, width 1.3 mm. Coloration usually uniformly yellowish, sometimes dark red-brown due to heavy sclerotization.Holotype with 48p+3a+T; paratype with 45p+4+T; paratypes with 40-59p+4-1a+T. Antennae much more elongate than usual (Figs 27A; 29A). Ocelli absent. Gnathochilarium with (n = 1) or without (n = 2) division of mentum (Fig. 29B).

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40ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. AB CD FIG. 28. — Glyphiulus basalis n. sp., paratype: A , caudal part of body, lateral view; B , same, ventral view; C , midbody segment section, frontal view; D , claw of a midbody leg. Scale bars: A, C, 0.5 mm; B, 0.2 mm; D, 0.05 mm.Crests on collum, dorsad of a well-developed lateral carina, nearly completely obliterated, pattern visible only as (4)5+m+5(4) extremely faint undulations at caudal margin (Fig. 27A, B). Carinotaxy of subsequent metaterga typical (Fig. 27D; 28D), but only two crests present below rather strikingly elongated ozoporiferous tubercles; all other crests distinctly underdevel-oped (Figs 27A, D; 28A, C). From segment 7 or 8 on, median crest clearly doubled anteriorly (Fig. 27D). Midbody segments round in cross-section (Fig. 28C). A low, but evident, transverse pleural ridge present behind gonopod opening on male segment 7. Epiproct narrower caudally, hypoproct more narrowly emarginate at caudal margin (Fig. 28B).Legs elongate (Figs 27C; 28C; 29F), longer than midbody height; claws strongly elongate, with a very small spine at base (Fig. 28D). Male legs 1 unusual in being fully leg-like, only very modestly reduced: a coxa, a 3-segmented telopodite and a claw supplied with a spine at base are all very clearly expressed (Fig. 29C); male legs 2 and 3 typical (Fig. 29D, E), but penes devoid of strong setae distolaterally.Anterior gonopods with evident, subunciform protuberances at median corners of coxosternum (Fig. 29G). Posterior gonopods with a ” agelliform and micropilose distal process (Fig. 29H). REMARKSBased on morphological evidence alone, this species seems to be a troglobite.

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41 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) A B C DE F G H FIG. 29. — Glyphiulus basalis n. sp., paratype: A , antenna; B , gnathochilarium, caudal view; C , legs 1, caudal view; D , leg 2, caudal view; E , leg 3; F , leg 11; G , anterior gonopods, caudal view; H , right posterior gonopod, caudal view. Scale bar: A-F, 0.4 mm; G, H, 0.3 mm.

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42ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. KEY TO SPECIES OF GLYPHIULUS GERVAIS, 1847 OF THE GRANULATUS-GROUP1. Carinotaxy patterns of collum and body segments typical, all crests well-developed: carinotaxy formula of collum: I-VI+7a+pc+ma+pc+7a+VI-I, i.e. middle part of collum with two abbreviated paramedian crests over front 2/3 collum, as well as three abbreviated crests medially in caudal 1/3 collum (Figs 2A; 10B; 13B; 15B; 17B); carinotaxy of midbody segments 3(2)+I+4+I+3(2) and 3(2)+i+3+i+3(2) (Figs 2B-F; 3B; 7C-F; 10C, D; 13C, D; 17C, D; 19A-C, E; 21; 22D; 27D), rarely even with a third row of 3(2)+i+3+i+3(2) (Fig. 15C, D), but median crest always doubled anteriorly. Relatively small species: length of adults 25 mm, width 1.3 mm ........................................................................... 2 „ Carinotaxy patterns of collum and body segments deviating from the typical ones, i.e. crests on collum either at least partly obliterated (Figs 7B; 19A, B; 21B; 24A, B; 27A, B) or, more seldom, showing a di erent formula, or some crests hypertrophied and differentiated, whereas midbody segments with either a typical (2+I+4+I+2 and 2+i+3+i+2) or an atypical formula, e.g., 3(2)+I+3+I+3(2) (Fig. 24). Often larger species .............. 7 2. Midbody segments in cross-section slightly compressed laterally (Fig. 13C). Legs slightly elongate, about as long as midbody height. Yunnan Prov., China ............................... 3„ Midbody segments round in cross-section (Figs 3B; 19E; 22D; 25B), not compressed laterally. Legs usually slightly shorter, rarely evidently longer, than midbody height .... 43. Antennae elongate (Fig. 12A). Spine at base of claw more than half as long as claw (Figs 11D; 12D, E) ................................................................. G. paragranulatus n. sp. „ Antennae stout (Fig. 14A). Spine at base of claw less than half as long as claw (Fig. 14D) ................................................................................................. G. semigranulatus n. sp. 4. Crests on midbody segments divided into three tubercles (Fig. 15D, E). Epiproct with a small but evident middorsal tubercle (Fig. 15E). Posterior gonopods with a plumose and sub” agelliform distal process (Fig. 16E, F) ........................................... G. beroni n. sp. „ Crests on midbody segments divided into two tubercles (Figs 2D; 7C, D; 17C, D; 21C, D). Epiproct without a middorsal tubercle (Figs 2E; 17E). Posterior gonopods with a plumose but subspatuliform distal process (Fig. 18H) ................................................ 55. Paraprocts evidently elevated, ridge-like along midline (Fig. 17F). Anterior gonopods with thicker telopodites, distomedial corners subunciform (Fig. 18G). Northern Laos .......... ........................................................................................................... G. bedosae n. sp.„ Paraprocts virtually ” at and smooth at midline (Figs 3C, D; 8B; 25D). Anterior gonopods with telopodites more slender, distomedial corners di erent ....................................... 66. Legs evidently shorter than midbody height (Fig. 3B). Antennae short and stout (Figs 3A; 4A). Anterior gonopods with characteristic, very high, distomedial processes (Figs 3E; 5A, B; 6C). Pantropical ........................................................................... G. granulatus„ Legs only slightly shorter than midbody height. Antennae slightly longer and more slender (more like in Figure 26A). Anterior gonopods much like in Figure 18G, distomedial processes low and broadly rounded. Hubei Prov., China ............................ G. lipsorum 7. Collum conspicuously hypertrophied due to crests I-IV, but paramedian crests complete and contrastingly low in anterior 1/3 collum. Vestigial 2-segmented leg remains on male sternum 1. Anterior gonopods much as in G. granulatus (Fig. 5A, B), distomedial processes similarly high but much wider. Posterior gonopods strongly constricted in distal 1/3. Mountains of central Vietnam .................................................................. G. capucinus „ Collum not hypertrophied, often some of its crests more or less obliterated. Male legs 1 either completely reduced to 1-segmented tubercular (usually) or leg-like. Anterior

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43 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) gonopods usually with much lower distomedial processes, if any. Posterior gonopods not so constricted in distal part (Fig. 7F, G) ...................................................................... 8 8. Midbody segments with median crest doubled anteriorly (Figs 7C; 19C; 21D; 27D). At least some crests on collum partly obliterated (Figs 7B; 21A, B; 24A, B), only seldom reduced very strongly (Fig. 27A, B) ............................................................................ 9„ Midbody segments with median crest not doubled anteriorly (Fig. 24C). Carinotaxy pattern of collum either atypical or di cult to trace due to crest reduction (Figs 19A, B; 24B) ......................................................................................................................... 14 9. Midbody segments slightly compressed laterally in cross-section .............................. 10 „ Midbody segments round in cross-section (Figs 19E; 28C) ...................................... 1110. Length of adults 19-28 mm, width 1.0-1.3 mm. Collum with 7+m+7 small rounded crests well visible at least near caudal edge (Figs 7B; 8A). Midbody segments with three lateral crests below ozoporiferous one (Fig. 7D, F). Claws as well as entire legs shorter (Figs 7D, F; 8C). Anterior gonopod telopodites especially slender, distomedial corners of coxosternum broadly rounded, lateral corners clearly dentiform (Fig. 9E). Distal process of posterior gonopod ” agelliform and plumose (Fig. 9F, G). Troglobite from Yunnan Prov., China ........................................................................................ G. subgranulatus n. sp.„ Length of adults 40-55 mm, width 2.0-2.3 mm. Collum with only a few undulations barely traceable near caudal edge (Fig. 19A, B). Body segments showing only two lateral crests below ozoporiferous one (Fig. 19A). Claws as well as entire legs longer (Fig. 19E, F). Anterior gonopod telopodites not particularly slender, distomedial corners of coxosternum subunciform (Fig. 20G). Distal process of posterior gonopod ” agelliform but bare (Fig. 20H). Troglobite from Hunan Prov., China .......................... G. deharvengi n. sp.11. Carinotaxy pattern of midbody segments typical, with three lateral crests below ozoporiferous one (Fig. 21). Crests on collum visible at both posterior and anterior margins (Fig. 21A, B). Obvious troglomorphic traits absent. Guizhou Prov., China ..................................... 12 „ Carinotaxy pattern of midbody segments also typical, but with two lateral crests below ozoporiferous one (Figs 24C; 25B; 27C; 28C). Crests on collum (4)5+m+5(4), hardly visible only at posterior margin. Troglomorphic traits very obvious .......................... 1312. Collum with 7+7 ” at crests at anterior and 5+m+5 similarly ” at crests at posterior margin, central part nearly smooth. Anterior gonopod coxosternum with a small subunciform projection at distomedial corner. Distal process of posterior gonopod blade-shaped, bare ......................................................................................................... G. quadrohamatus „ Collum with 7+m+7 ” at crests at posterior margin, anteromedial part nearly smooth, but carinotaxy pattern discernible as typical (Fig. 21A, B). Anterior gonopod coxosternum particularly simple, subtruncate (Fig. 23F). Distal process of posterior gonopod ” agelliform and plumose (Fig. 23G, H) ....................................................... G. pergranulatus n. sp.13. Male legs 1 leg-like (Fig. 29C). Ozoporiferous tubercles unusually prominent compared with the other, strongly reduced crests. Troglobite from Sichuan Prov., China ... G. basalis n. sp. „ Male legs 1 strongly reduced (as in Figure 4D). Ozoporiferous tubercles low, much wider than high ................................................................................................................. 1414. Collum with rather evident crests at caudal margin, but entirely smooth over anterior 1/3. Ocelli completely reduced. Troglobite from Guangxi Prov., China ..... G. anophthalmus „ Collum with crests barely visible in posterior 1/3, being totally obliterated over anterior 2/3. Ocelli present but unpigmented, reduced in number to about 6. Troglobite from Guizhou Prov., China .................................................................................... G. balazsi

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44ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. 15. Carinotaxy pattern of midbody segments strongly atypical, formula being 2+I+2+I+2 and 2+i+3+i+2. Collum with 5+ma+5 ” at crests/undulations barely visible at posterior margin. Guangxi Prov., China .............................................................. G. adeloglyphus„ Carinotaxy pattern of midbody segments easily derivable from typical one. Collum varying from strongly carinate to virtually smooth, usually easily derived from typical carinotaxy pattern ..................................................................................................................... 16 16. Collum strongly carinate, no crests being obliterated. Legs and claws relatively short, legs shorter than midbody height .................................................................................... 17„ Collum evidently carinate only at posterior margin at most, crests always being obliterated over central and/or anterior parts (Fig. 24A, B), or collum entirely smooth. Presumed troglobites, legs at least as long as midbody height (Fig. 25B); claw slender and long (Fig. 25D), at least as long as tarsus .......................................................................... 1817. Length of adults 17-30 mm, width up to 1.2 mm. Carinotaxy pattern of collum deviating from typical mainly in having both paramedian crests complete, holoseriate, with formula I-VI+7a+P+ma+P+7a+VI-I. Epiproct with three weak crests dorsally. Anterior gonopod coxosternum with a short distomedian outgrowth. Guangxi Prov., China (also introduced to Okinawa Island, Japan) .................................................................. G. septentrionalis„ Length of adults up to 85 mm, width up to 5.0 mm. Carinotaxy pattern of collum deviating in having at least both paramedian and median crests incomplete (formula I+2c+III…IV+(5c?)+VI+7p+8a+pc+ma+pc+8a+7p+VI+(5c?)+IV-III+2c+I). Epiproct with a weak crest dorsally. Anterior gonopod coxosternum much as in G. granulatus , with a very long and slender distomedian process. Mountains of central Vietnam .... G. superbus 18. Dorsal crests on midbody segments high, clearly divided into two halves. Ozoporiferous tubercles higher than wide. Guangxi Prov., China ................................ G. melanoporus„ Dorsal crests on midbody segments invariably very low and undivided (Fig. 24A, D). Ozoporiferous tubercles wider than high (Fig. 24C). Guizhou Prov., China ............. 19 19. Collum completely smooth. Hypoproct very poorly emarginate medially .... G. rayrouchi„ Collum with poor but evident crests/undulations, a rather typical carinotaxy pattern visible at least medially (Fig. 24A, B). Hypoproct emargination nearly notch-like caudomedially (Fig. 25A) ............................................................................................ G. latellai n. sp.CONCLUDING REMARKSNow that the granulatus -group contains 20 spe-cies, and numerous presumed congeners from other groups that will be described elsewhere, it seems appropriate to provide brief analyses of variation in Glyphiulus and of the distribution of its constituent species. For the sake of convenience, and in agreement with Verhoe (1936) and Loksa (1960), it seems best for the moment to accept the granulatus -group as being the same as Glyphiulus s.s. However, with further accumulation of material and its analysis, it may become necessary to adopt a broader concept of Glyphiulus .Although the value of the classical division of the gnathochilarial mentum into a promentum and a eumentum has long been disputed and suspected as being overestimated in the systematics of camba-loid Juliformia (Verhoe 1936; Attems 1938), the present discovery of infraspeci“ c variation in this character in three species of the granulatus -group, G. granulatus , G. subgranulatus n. sp. and G. basalis n. sp. , was quite surprising. is means that, within the Cambalopsidae at least, the mentum only tends to be divided in Glyphiulus. Similarly, the di erent degrees of setation of the gnathochilarium, from oligoto polytrichous, also seem to re” ect noth-ing but individual variation. In other words, the

PAGE 39

45 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) 12 13 5 1, 2, 3 4 2 11 15 17, 18 19 10 9 6 7 8 16 14 Sichuan Yunnan MYANMAR Guizhou Hubei Hunan VIETNAM LAOS THAILAND CAMBODIA 400 km FIG. 30. — Distribution of species Glyphiulus Gervais, 1847 of the granulatus -group (that of G. granulatus (Gervais, 1847) being pantropical, it is omitted here), shown more or less from west to east: 1 , G. paragranulatus n. sp.; 2 , G. semigranulatus n. sp.; 3 , G. beroni n. sp.; 4 , G. subgranulatus n. sp.; 5 , G. bedosae n. sp.; 6 , G. pergranulatus n. sp.; 7 , G. rayrouchi Mauriès & Nguyen Duy-Jacquemin, 1997; 8 , G. latellai n. sp.; 9 , G. quadrohamatus Chen & Meng, 1991; 10 , G. balazsi (Loksa, 1960); 11 , G. anophthalmus (Loksa, 1960); 12 , G. capucinus Attems, 1938; 13 , G. superbus Silvestri, 1923; 14 , G. basalis n. sp.; 15 , G. lipsorum Mauriès & Nguyen Duy-Jacquemin, 1997; 16 , G. deharvengi n. sp.; 17 , G. melanoporus Mauriès & Nguyen Duy-Jacquemin, 1997; 18 , G. septentrionalis Murakami, 1975; 19 , G. adeloglyphus Zhang & Li, 1982.structure of the gnathochilarium seems more or less useless at the speci“ c level. e division versus non-division of the mentum should only be applied to the genus level and even higher classi“ cation of cambaloids with extreme care, representing more of an evolutionary trend to be checked using suf“ ciently representative material. As regards the carinotaxy in Glyphiulus , the patterns, but not the degrees of expression, of crests on the collum and body segments seem to be quite stable. Hence the patterns prove to be highly useful in the classi“ cation, often allowing the grouping of species. e degrees of expression of the crests prove to be very stable both between sexes and at

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46ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. various developmental stages, making it a very use-ful character for species discrimination. e carinotaxy pattern of the collum is often, but not always, correlated to that of the metaterga. In several cases at least, this correlation is di cult to elucidate because the crests on the collum are nearly or entirely obliterated. Crest reduction always concerns the anterior and/or central parts of the collum. Complete loss of crests/undulations is rare (i.e. G. rayrouchi ). In general, crest arrange-ments and forms tend to be more variable on the collum than on the metaterga. Crest size reduc-tion or hypertrophy concerns all of the metatergal crests simultaneously or nearly so, except perhaps the ozoporiferous tubercles, which always remain the largest and sometimes (in G. basalis n. sp.) even become enlarged against a background of the other strongly reduced crests. Reduction is especially evi-dent due to loss of the ventrolateral crest, which turns into a stria similar to several other striae located further ventrad.Changes in the carinotaxy formulae of the collum from the typical I-VI+7a+pc+ma+pc+7a+VI-I down to I-IV(V)+5(6)a+pc+ma+pc+5(6)a+(V)IV-I, due to fusion/loss of one or two lateralmost crests, are rather common. e pattern shown by the paramedian and median crests, when these are traceable, seems to be much more stable and important. Usually it is pc+ma+pc, only seldom P+ma+P ( G. septentrionalis ). Only one species, G. capucinus , has the crests strongly di erentiated (some are hypertrophied, some others reduced), but the formula is only slightly di erent from that of G. superbus .Variation in the carinotaxy formulae of the meta-terga is more modest. Usually the crests are clearly divided near the middle into two half-crests, thus forming two transverse rows of tubercles. Usually the tubercles in the anterior row are smaller than those in the posterior row. In one case only ( G. beroni n. sp.), most of the crests are divided transversely into three rows, whereas the crests are undivided only in G. rayrouchi and G. latellai n. sp. A typical pattern is 3(2)+I+4+I+3(2) and 3(2)+i+3+i+3(2), meaning that the median crest is doubled anteriorly. is formula is highly characteristic of the granulatus -group, with only relatively few, rather minor deviations that show no anterior duplication of the middorsal crest. e strongest deviation observed is 2+I+2+I+2 and 2+i+3+i+2 ( G. adeloglyphus ).Several species of the group are presumed troglobites. Besides showing the typical traits of troglomorphism (general depigmentation, eye reduction, elongation of legs, claws and anten-nae, gigantismŽ, although all relatively modestly expressed in comparison with some examples in Polydesmida or even Julida), such species tend to have more or less strongly obliterated crests on the collum, as well as reduced crests on the metaterga. In addition, they tend to show a more narrowly rounded epiproct and a more narrowly emarginate hypoproct. Although there is a pat-tern of elongated claws in the troglobitic species of Glyphiulus s.s., there seems to be no correlation between this and the size of the accessory spine at the claw base.Legs 1 of the male are normally strongly reduced, but in G. superbus , besides a small coxa, a rudimentary telopodite segment is still discernible. Furthermore, G. basalis n. sp. seems to be even more primitive in this respect, showing a nearly fully developed, 3-segmented telopodite crowned with a typical claw and a spine at its base. What remains invariable is that the sternum is devoid of any median or paramedian structures, being supplied instead with two widely separated prongs curved caudally. is character is of primary importance in delimiting Glyphiulus s.s. from the other cambalopsids. Because the vulvae in Glyphiulus are surprisingly monotonous in structure, they are hardly of use systematically. In addition, the peripheral charac-ters of the female (and even of the juveniles) are so abundant and evident that they are quite suf-“ cient for species recognition, as well as safe sex and stadium matches. Unsurprisingly, there are many species of Cambalopsidae described solely from female material. e general distribution of the granulatus -group shows quite a coherent pattern (Fig. 30), occupy-ing a region between Sichuan, Hubei and Shanghai in the North and central Vietnam in the South. ere is no doubt that these generally subtropical to tropical, highly montane and often karstic areas contain many new species of Glyphiulus waiting to be found and described.

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47 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) e fact that the species of the granulatus -group are mostly highly local in distribution deserves attention. is can often be accounted for by their being troglobites naturally restricted to their cave or a group of caves. However, the impression that most of the species are scarcely more than troglophiles suggests that a wealthy, barely sampled fauna exists epigeically as well. is is because most of the expe-ditions to these often remote and poorly accessible places have been speleological, concentrating on the mystiqueŽ of the underground world, thus giving the biased impression that cambalopsids in general, and Glyphiulus species in particular, are especially common in caves. Above-ground explorations will certainly reveal a far richer fauna of forest-dwelling species in that region. Summarising the poor state of current knowledge, we seem to have only touched the top of an iceberg. Acknowledgements is work only became possible through the support provided to the “ rst author by the MNHN. Anne Bedos and Louis Deharveng (both MNHN), Josiane and Bernard Lips (Villeurbanne, France), Petar Beron (NMNHS) and Leonardo Latella (MCSNV) are deeply thanked for the precious material they o ered for study and for donating it entirely or partly to MNHN. Mark Judson (MNHN) kindly corrected the English text. Both Richard L. Ho man (Virginia Museum of Natural History, Martinsville) and Henrik Engho (Natural History Museum of Denmark, Copenha-gen) provided constructive criticism of advanced drafts. Figure 1 was skilfully executed by Mrs Nadine E. Van Noppen (MRAC), while the previously unpublished Figures 6E and G are from Henri Brolemanns IconographieŽ kept at MNHN. REFERENCESATTEMS C. 1900. „ Dr. Brauers Myriopoden-Aus-beute auf den Seychellen im Jahre 1895. Zoologische Jahrbücher, Abteilung für Systematik, Geographie und Biologie der Tiere 13: 133-171. ATTEMS C. 1926. „ Myriapoda in KÜKENTHAL W. & KRUMBACH T. (eds), Handbuch der Zoologie , IV, 1, Progoneata, Chilopoda, Insecta . W. de Gruyter & Co., Berlin; Leipzig: 1-402.ATTEMS C. 1938. „ Die von Dr C. Dawydoff in Französisch Indochina gesammelten Myriopoden. Mémoires du Muséum national dHistoire naturelle n.s. 6 (2): 187-353.ATTEMS C. 1953. „ Myriopoden von Indochina. Gesam-melt von Dr C. Dawydo (1938-1939). Mémoires du Muséum national dHistoire naturelle sér. A (Zoologie), 5 (3): 133-230. BROLEMANN H. W. 1931. „ Myriapodes recueillis par Mme Pruvot en Nouvelle-Calédonie et aux Loyalty. Archives de Zoologie expérimentale et générale 72 (2): 275-316.CARL J. 1911. „ Drei neue Diplopoden des Genfer Museums. Revue suisse de Zoologie 19: 397-407. CHEN J. X. & MENG W. X. 1991. „ A new species of the genus Glyphiulus (Diplopoda, Spirostreptida, Cambalopsidae). Acta Zootaxonomica Sinica 16: 394-397 (in Chinese, English summary).CROSLAND M. W. J. 1994. „ Millipede myrmecophile, Glyphiulus granulatus , found with the primitive ant Harpegnathos venator . Journal of Entomological Science 29: 141-142. ENGHOFF H. 1993. „ Haplopodous diplopods: a new type of millipede body construction discovered in cambalopsid juveniles (Diplopoda, Spirostreptida). Acta Zoologica 74 (3): 257-261.ENGHOFF H. 2005. „ The millipedes of Thailand (Diplopoda). Steenstrupia 29: 87-103.ENGHOFF H., DOHLE W. & BLOWER J. G. 1993. „ Ana-morphosis in millipedes (Diplopoda) … the present state of knowledge with some developmental and phylogenetic considerations. Zoological Journal of the Linnean Society 109: 103-234. ENGHOFF H., GOLOVATCH S. I. & NGUYEN DUC ANH 2004. „ A review of the millipede fauna of Vietnam (Diplopoda). Arthropoda Selecta 13 (1-2): 29-43.GEOFFROY J.-J. & GOLOVATCH S. I. 2004. „ Some polydesmidan millipedes from caves in southern China (Diplopoda: Polydesmida), with descriptions of four new species. Arthropoda Selecta 13 (1-2): 19-28.GERVAIS P. 1847. „ Myriapodes, in WALCKENAER C. A. & GERVAIS P. (eds), Histoire naturelle des Insectes Aptères , Tome 4e. Librairie encyclopédique de Roret, Paris: 1-623.GOLOVATCH S. I., GEOFFROY J.-J. & MAURIÈS J.-P. 2006a. „ Four new Chordeumatida (Diplopoda) from caves in China. Zoosystema 28 (1): 75-92.GOLOVATCH S. I., GEOFFROY J.-J. & MAURIÈS J.-P. 2006b. „ Review of the millipede genus Hyleoglomeris Verhoe , 1910 (Diplopoda, Glomerida, Glomeridae), with descriptions of new species from caves in Southeast Asia. Zoosystema 28 (4): 887-915. GOLOVATCH S. I., GEOFFROY J.-J. MAURIÈS J.-P. & VAN DEN SPIEGEL D. in press. — Review of the millipede

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48ZOOSYSTEMA • 2007 • 29 (1)Golovatch S. I. et al. genus Glyphiulus Gervais, 1847, with descriptions of new species from Southeast Asia (Diplopoda, Spirostreptida, Cambalopsidae). Part 2: the javanicus -group. Zoosystema .HOFFMAN R. L. 1977. „ Diplopoda from Malayan caves collected by M. Pierre Strinati. Revue suisse de Zoologie 84 (3): 699-719.HOFFMAN R. L. 1980. „ Classi“ cation of the Diplopoda . Muséum dhistoire naturelle, Genève, 237 p. (dated 1979, published 1980).JEEKEL C. A. W. 1963. „ Diplopoda of Guiana (1-5), in GIJSKES D. C. & WAGENAAR HUMMELINCK P. (eds), Studies on the Fauna of Suriname and Other Guyanas 4. Martinus Nijho , e Hague: 1-157.JEEKEL C. A. W. 1971. „ Nomenclator generum et familiarum Diplopodorum. A list of the genus and family-group names in the class Diplopoda from the 10th edition of Linnaeus, 1758, to the end of 1957. Monogra“ eën van de Nederlandse Entomologische Vereiniging 5: i-xii + 1-412 (dated 1970, published 1971).JEEKEL C. A. W. 1985. „ e distribution of the Diplo-cheta and the lostŽ continent Paci“ ca (Diplopoda). Bijdragen tot de Dierkunde 55 (1): 100-112.JEEKEL C. A. W. 2004. „ A bibliographic catalogue of the CambaloideaŽ (Diplopoda, Spirostreptida). Myriapod Memoranda 7: 43-109.KARSCH F. 1881. „ Neue Juliden des Berliner Museums, als Prodromus einer Juliden-Monographie. Zeitschrift für die gesammten Naturwissenschaften 54: 1-79.KORS"S Z. 2004. „ Checklist and bibliography of millipedes (Diplopoda) of Taiwan. Collection and Research (NMNS, Taichung) 17: 11-32.KRABBE E. 1982. „ Systematik der Spirostreptidae (Diplopoda, Spirostreptomorpha). Abhandlungen des Naturwissenschaftlichen Vereins in Hamburg , Neue Folge 24: 1-476.LOKSA I. 1960. „ Einige neue Diplopodenund Chilopodenarten aus chinesischen Höhlen. Acta Zoologica Academiae Scientiarum Hungaricae 6 (1-2): 137-148.MAURIÈS J.-P. 1970. „ Examen des types des genres Cambalomorpha et Cambalopsis Pocock, 1895. Essai de classi“ cation des Glyphiulinae Verhoe , 1936 (Diplopoda, Cambalidea). Bulletin du Muséum national dHistoire naturelle sér. 2, 42 (3): 509-519.MAURIÈS J.-P. 1977. „ Le genre Glyphiulus Gervais, 1847, et sa place dans la classi“ cation des Cambalides, à propos de la description dune nouvelle espèce du Viêt-Nam (Diplopoda, Iulida, Cambalidea). Bulletin du Muséum national dHistoire naturelle sér. 3, 431 (Zoologie 301): 243-250. MAURIÈS J.-P. 1981. „ Craspedosomida, Stemmiulida et Cambalida (Myriapoda: Diplopoda) de Sri Lanka (Ceylan). Entomologica Scandinavica , Supplementum 11: 33-62.MAURIÈS J.-P. 1983. „ Cambalides nouveaux et peu connus dAsie, dAmérique et dOcéanie. I. Cambalidae et Cambalopsidae (Myriapoda, Diplopoda). Bulletin du Muséum national dHistoire naturelle 4e sér., sect. A, 5 (1): 247-276.MAURIÈS J.-P. 1987. „ Cambalides nouveaux et peu connus dAsie, dAmérique et dOcéanie. II. Pseudo-nannolenidae, Choctellidae (Myriapoda, Diplopoda). Bulletin du Muséum national dHistoire naturelle 4e sér., sect. A, 9 (1): 169-199.MAURIÈS J.-P. 1992. „ Sur la vraie place du genre Protosilvestria Handschin dans la classi“ cation des Diplopodes Iuliformes, in MEYER E., THALER K. & SCHEDL W. (eds), Proceedings of the 8th International Congress of Myriapodology. Berichte des Naturwis-senschaftlich-Medizinischen Vereins in Innsbruck, Supplementum 10: 23-31.MAURIÈS J.-P. 2005. „ Guizhousoma latellai gen. n., sp. n., de Chine continentale, type dune nouvelle famille de la superfamille des Neoatractosomatoidea (Diplopoda: Chordeumatida). Arthropoda Selecta 14 (1): 11-17.MAURIÈS J.-P. & NGUYEN DUY-JACQUEMIN M. 1997. „ Nouveaux Craspedosomides et Glyphiulides cavernicoles de Chine (Diplopoda). Mémoires de Biospéologie 24: 46-62.MURAKAMI Y. 1975. „ e cave myriapods of the Ryukyu Islands (I). Bulletin of the National Science Museum Series A (Zoology), 1 (2): 85-113.PETERS W. 1864. „ Eine Übersicht der im Königl. zoologischen Museum be“ ndlichen Myriopoden aus der Familie der Polydesmi, so wie Beschreibungen einer neuen Gattung, Trachyjulus , der Juli und neuer Arten der Gattung Siphonophora . Monatsbericht der Königlichen Preussischen Akademie der Wissenschaften zu Berlin 1864: 529-551. POCOCK R. I. 1893. „ Report upon the Myriopoda of the ChallengerŽ Expedition, with remarks upon the fauna of Bermuda. Annals and Magazine of Natural History (6) 11: 121-142.POCOCK R. I. 1894. „ Chilopoda, Symphyla and Diplopoda from the Malay Archipelago, in WEBER M. (ed.), Zoologische Ergebnisse einer Reise in Niederländisch Ost-Indien 3: 307-404.POCOCK R. I. 1895. „ Report upon the Chilopoda and Diplopoda obtained by P. W. Basset-Smith, Esq., Surgeon R. N., during the cruise in the Chinese seas of H.M.S. PenguinŽ, Commander W. U. Moore commanding. 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49 Review of Glyphiulus : the granulatus -group (Diplopoda, Cambalopsidae)ZOOSYSTEMA • 2007 • 29 (1) Yunnan, China, with remarks on the reduction of the second female leg-pair (Diplopoda: Callipodida: Caspiopetalidae). Journal of Natural History 39: 1875-1891.STOEV P. & GEOFFROY J.-J. 2004. „ Review of the millipede family Paracortinidae Wang & Zhang, 1993 (Diplopoda: Callipodida). Acta Arachnologica 53 (2): 93-103. VERHOEFF K. W. 1924. „ Results of Dr. E. Mjöbergs scienti“ c expeditions to Australia 1910-1913. 34. Myriapoda: Diplopoda. Arkiv för Zoologi 16 (5): 1-142.VERHOEFF K. W. 1936. „ Zur Kenntnis der Glyphiuliden (Cambaloidea). 143. Diplopoden-Aufsatz. Zoologischer Anzeiger 113 (3/4): 49-62.ZHANG C. Z. & LI Z. Y. 1982. „ Die Gattung Glyphiulus (Diplopoda: Cambalidea) von China. Sinozoologia 2: 85-93 (in Chinese, German summary). Submitted on 23 November 2005; accepted on 2 May 2006.


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