A Checklist of the Millipedes (Diplopoda) of Cambodia

Zootaxa 3973 (1): 175–184 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2015 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3973.1.7 http://zoobank.org/urn:lsid:zoobank.org:pub:60AB9881-B748-4A32-8800-5EEB489EE535 A checklist of the millipedes (Diplopoda) of Cambodia

NATDANAI LIKHITRAKARN1, SERGEI I. GOLOVATCH2,4 & SOMSAK PANHA3,4 1Division of Plant Protection, Faculty of Agricultural Production, Maejo University, Chiang Mai 50290, Thailand 2Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky pr. 33, Moscow 119071, Russia 3Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand 4Corresponding authors. E-mail: Somsak Panha ([email protected]); Sergei I. Golovatch ([email protected])

Abstract

At the present, the millipede fauna of Cambodia comprises only 19 species from 15 genera, 12 families and 8 orders. These counts certainly represent but a minor fraction of the country’s real diversity of Diplopoda even at the ordinal level, let alone at lower ones. Based on the available information from the adjacent parts of China, Thailand, Myanmar, Vietnam and/or Laos, the orders Glomerida, Platydesmida, Polyzoniida, Callipodida and Chordeumatida must occur in Cambodia, maybe also Stemmiulida and Siphonocryptida, but none has been recorded there yet. This shows that a lot more collecting effort is required to amass a representative material of Diplopoda of Cambodia to make it available for study.

Key words: millipede, taxonomy, fauna, Cambodia

Introduction

The Kingdom of Cambodia is a tropical country found on the peninsula of mainland Southeast Asia adjacent to the gulf of Thailand with a land area of 181,035 km2. Cambodia has a coastline of 435 km, and its land border of 2,438 km runs along Thailand to the west, Vietnam to the east and Laos PDR to the north (Map 1) (Kingdom of Cambodia 2010). The climate of Cambodia is tropical, dominated by the annual monsoon cycle, which is accompanied by alternating wet and dry seasons. Cambodia is considered a high forest cover country: in 2006, 59% of the country was covered by forest (Technical Working Group on Forestry & Environment 2007). The country also contains the largest freshwater lake in Southeast Asia, the Tonle Sap Lake. In terms of biodiversity, the country is believed to have over 130 mammal and more than 500 bird species. Some 300 species of freshwater fish have been identified, of which 215 are in the Tonle Sap (ADB 2000). Studies on the millipedes (Diplopoda) in Cambodia started with descriptions or records of 13 species contained in the works of Attems (1938, 1953), based on the collections made by staff of the Paris Museum during the 1930s in French Indochina. These two papers still remain the cornerstones of our knowledge of the Myriapoda of Indochina east of Thailand (Likhitrakarn et al. 2014a). As a result, at the present the fauna contains at least 19 species, one of them (Glomeridesmus sp.) yet requiring a closer identification. Cambodia remains the only country of Indochina which still lacks a comprehensive millipede checklist (cf. Enghoff et al. 2004; Enghoff 2005; Likhitrakarn et al. 2014a). To promote further studies on the Diplopoda of that country, as well as to pinpoint the existing shortcomings, the following catalogue is provided. A complete bibliography list relevant to the Cambodian fauna is also compiled.

Material and methods

The present paper is a conventional checklist restricted to the Diplopoda documented from Cambodia. In the catalogue sections, D stands for the original description, subsequent descriptive notes or appearance in a key, R for a subsequent record or records and M for a mere mention. A reference to the original description is always given,

Accepted by W. Shear: 26 Mar. 2015; published: 16 Jun. 2015 175 regardless if this concerns the fauna of Cambodia or not. The new material collected by members of the Animal Systematics Research Unit, Chulalongkorn University in February 2015 is now housed in the Museum of Zoology, Chulalongkorn University (CUMZ), Bangkok, Thailand. The French or German locality names quoted in Attems (1938, 1953) and Demange (1961) can be updated in the following tabular form: Angkor = Siem Reap Province, Angkor, 13°22′0.84″N, 103°51′46.8″E Angkor Vat = Siem Reap Province, Angkor Vat, 13°24′45″N, 103°52′0″E Berg Cardamon = Koh Kong Province, Cardamon Mountains Bokoz, Bokkor = Kampot Province, Phnom Bokor Kampong Speu = Kampong Speu Province, Kampong Speu Kampot = Kampot Province, Kampot Kratiée (Makong) = Kratié Province, Kratié Massif de l’Eléphant (Elephant Mountain) = Kampot Province, Dâmrei Mountains Réam = Sihanoukville Province, Ream Sre Umbell = Koh Kong Province, Sre Ambel

MAP. Distribution of millipedes in Cambodia (18 species): 1 National highway 5 bridge near Serei Sophorn River. 2 Angkor Vat. 3 Angkor. 4 Kratié. 5 Cardamon Mountains. 6 Kampong Speu. 7 Kien Krol G Cave. 8 Sre Ambel. 9 Phnom Bokor. 10 Ream. 11 Dâmrei Mountains. 12 near Kampong Trach. 13 Southeast part of Koh Tang.

Order Polyxenida

Family Polyxenidae

Genus Alloproctoides Marquet & Condé, 1950

1. Alloproctoides dawydoffi (Attems, 1938) Monographis kraepelini dawydoffi Attems, 1938: 321 (D). Monographis kraepelini dawydoffi—Attems, 1953: 153 (R). Alloproctoides dawydoffi Nguyen Duy-Jacquemin & Condé, 1967: 55 (D), 1982: 109 (D); Golovatch, 1983: 185 (M); Nguyen Duy-Jacquemin & Geoffroy, 2003: 97 (M); Enghoff et al., 2004: 30 (R). Record from Cambodia: Kampot Province, Phnom Bokor (Attems 1953). Also known from Vietnam (Attems 1938) and Atoll Tizard (= Spratly Archipelago) (Attems 1953).

Order Sphaerotheriida

Family Zephroniidae

Genus Zephronia Gray, 1832

2. Zephronia dawydoffi Attems, 1953 Zephronia dawydoffi Attems, 1953: 156 (D). Zephronia dawydoffi—Jeekel, 2001a: 18 (R). Zephronia cambodjana Attems, 1953: Tafel III (D). Records from Cambodia: Sihanoukville Province, Ream; Koh Kong Province, Sre Ambel; Kampot Province, Kampot; Kampong Speu Province, Kampong Speu (Attems 1953). Endemic to Cambodia. Remark. The figure caption to Zephronia dawydoffi in Attems (1953) contains a misprint reading Zephronia cambodjana.

Order Glomeridesmida

Family Glomeridesmidae

Genus Glomeridesmus Gervais, 1844

3. Glomeridesmus sp. Glomeridesmus sp.—Shelley, 2011: 3 (M). Record from Cambodia: Kampot Province, Phnom Laang, Kien Krol G Cave (Shelley 2011). Remark. Unidentified Glomeridesmus species have also been reported from Eastern Malaysia, Island of New Guinea, Papua New Guinea, Oceania, Thailand, Philippines and Indonesia (Shelley 2011).

Order Siphonophorida

Family Siphonorhinidae

Genus Siphonorhinus Pocock, 1984

MILLIPEDES OF CAMBODIA Zootaxa 3973 (1) © 2015 Magnolia Press · 177 4. Siphonorhinus coniceps (Attems, 1936) Siphonophora coniceps Attems, 1936: 314 (D). Indiozonium coniceps—Verhoeff, 1941: 215 (M). Siphonorhinus coniceps—Carl, 1941a: 573 (M); Jeekel, 2001b: 47 (R). Pterozonium coniceps—Attems, 1951: 231 (M), 1953: 198 (D, R). Zinaceps coniceps—Chamberlin & Wang, 1953: 13 (M). Record from Cambodia: Koh Kong Province, Sre Ambel (Attems 1953). Also known from India (Attems 1936).

Family Siphonophoridae

Genus Siphonacme Cook & Loomis, 1928

5. Siphonacme dawydoffi (Attems, 1938) Siphonophora dawydoffi Attems, 1938: 311 (D). Annamnium dawydoffi—Verhoeff, 1941: 217 (D); Golovatch, 1983: 185 (M). Siphonacme dawydoffi—Attems, 1951: 226 (M); 1953: 198 (R); Jeekel, 2001b: 53 (R); Enghoff et al., 2004: 33 (R). Record from Cambodia: Kampot Province, Phnom Bokor (Attems 1938). Also known from Vietnam (Attems 1938) and Laos (Attems 1953).

Order Julida

Family Julidae

Genus Nepalmatoiulus Mauriès, 1983

6. Nepalmatoiulus crassus (Attems, 1938) Fusiulus crassus Attems, 1938: 251 (D). Fusiulus crassus—Attems, 1953: 195 (R). “Anaulaciulus” crassus Golovatch, 1983: 182 (M). Nepalmatoiulus crassus Enghoff, 1987: 304 (D); Enghoff et al., 2004: 33 (R). Record from Cambodia: Kratié Province, Kratié (Attems 1953). Also known from Vietnam and Laos (Attems 1938, 1953). Remarks. This species, also reported from Pic de Lang Biang (2,400 m a.s.l.), Champasak Province, Xieng Kuang (1,500 m a.s.l.), Xiang Khouang Province and Lung Prabang, Luang Prabang Province, Laos (Attems 1953), was inadvertently omitted from the diplopod checklist of Laos by Likhitrakarn et al. (2014a).

Order Spirobolida

Family Pachybolidae

Genus Aulacobolus Pocock, 1903

7. Aulacobolus rubropunctatus Attems, 1938 Aulacobolus rubropunctatus Attems, 1938: 261 (D). Atopochetus rubrodorsalis Attems, 1953: 191 (D). Aulacobolus rubropunctatus—Hoffman, 1962: 777 (D); Jeekel, 2001c: 50 (R). Atopochetus rubrodorsalis—Jeekel, 1971: 193 (M). Records from Cambodia: Sihanoukville Province, Ream (Attems 1938, 1953); Koh Kong Province, Sre Ambel and

Family Trigoniulidae

Genus Trigoniulus Pocock, 1894

8. Trigoniulus corallinus (Gervais, 1847) Iulus corallinus Gervais, 1847: 275 (D). Trigoniulus corallinus Hoffman, 1994: 20 (M). Trigoniulus corallinus—Jeekel, 2001c: 72 (R); Enghoff et al., 2004: 35 (R); Enghoff, 2005: 90 (R). New material: 2 males, 2 females (CUMZ), Cambodia, Banteay Meanchey Province, Sisophon, National highway 5 bridge near Serei Sophorn River, 13°24′48.2″N, 102°55′26.1″E, 22 m a.s.l., 06.02.2015, leg. C. Sutcharit, W. Siriwut, P. Jirapatrasilp and R. Srisonchai. This is a synanthropic species also known from numerous tropical islands throughout the world, as well as from coastal and continental sites (e.g. Jeekel 2001c; Enghoff et al. 2004; Enghoff 2005).

Order Spirostreptida

Family Cambalopsidae

Genus Glyphiulus Gervais, 1847

9. Glyphiulus superbus Silvestri, 1923 Glyphiulus superbus Silvestri, 1923: 190 (D). Glyphiulus superbus—Attems, 1936: 251 (R); 1938: 266 (D); 1953: 195 (R); Mauriés, 1970: 518 (M); 1977: 246 (D); Jeekel, 1971: 111 (M), 2004: 54 (R); Golovatch, 1983: 183 (M); Enghoff et al., 2004: 35 (R); Golovatch et al., 2007: 11 (R). Koinoglyphius superbus—Carl, 1941b: 287 (D). Record from Cambodia: Kampot Province, Kampot (Attems 1953). Also known from Vietnam (Silvestri 1923).

Genus Plusioglyphiulus Silvestri, 1923

10. Plusioglyphiulus dubius (Attems, 1938) Glyphiulus dubius Attems, 1938: 272 (D). Plusioglyphiulus dubius—Mauriès, 1970: 510 (M), 1983: 272 (D); Hoffman, 1977: 175 (M); Jeekel, 2004: 57 (R); Golovatch et al., 2009: 74 (R). Record from Cambodia: Kampot Province, Phnom Bokor (Attems 1938). Endemic to Cambodia.

11. Plusioglyphiulus boutini Mauries, 1970 Plusioglyphiulus boutini Mauriès, 1970: 509 (D). Plusioglyphiulus boutini—Hoffman, 1977: 175 (M); Mauriès, 1983: 272 (D); Boutin, 2001: 1760 (M); Golovatch et al., 2009: 74 (R). Record from Cambodia: Kampot Province, near Kampong Trach (Mauriès 1970), 10.554° N, 104.471° E (Golovatch et al. 2009). Endemic to Cambodia.

Genus Rhynchoproctus Pocock, 1894

12. Rhynchoproctus proboscideus Pocock, 1894 Rhynchoproctus proboscideus Pocock, 1894: 386 (D). Rhynchoproctus proboscideus—Sinclair, 1901: 522 (R); Attems, 1914: 289 (R); 1942: 80 (R); Chamberlin, 1920: 173 (R); Demange, 1961: 245 (R); 1986: 852 (R); Wang & Tang, 1965: 430 (D); Jeekel, 1971: 135 (M); 2006: 40 (R); Pimvichai et al., 2010: 70 (R). Rhynchoproctus minor Silvestri, 1897: 2 (D). Rhynchoproctus minor—Demange, 1961: 245 (M); 1969: 258 (D); Jeekel, 2006: 40 (R). Rhynchoproctus longipes Silvestri, 1897: 2 (D). Rhynchoproctus longipes—Demange, 1961: 245 (M); 1969: 256 (D); Jeekel, 2006: 40 (R). Cambodjostreptus castaneus Attems, 1953: 194 (D). Cambodjostreptus castaneus—Demange, 1983 (M); Jeekel, 1971: 122 (M); 2006: 41 (R). Records from Cambodia: Koh Kong Province, Cardamon Mountains and Kampot Province, Dâmrei Mountains, 1,080 m a.s.l. (Attems 1953). Also known from Thailand, Malaysia and Indonesia (Jeekel 2006; Pimvichai et al. 2010).

Genus Thyropygus Pocock, 1894

13. Thyropygus resimus Attems, 1938 Thyropygus resimus Attems, 1938: 283 (D). Thyropisthus resimus—Attems, 1942: 78 (M); Demange, 1961: 142 (D, R). Thyropygus allevatus resimus—Hoffman, 1975: 136 (D); Jeekel, 2006: 13 (R). Thyropygus resimus—Pimvichai et al., 2011a: 53 (D, R). Record from Cambodia: Siem Reap Province, Angkor Vat (Demange 1961). Also known from Laos (Attems 1953; Pimvichai et al. 2011a), Thailand and Myanmar (Pimvichai et al. 2011a).

14. Thyropygus cuisinieri Carl, 1917 Thyropygus cuisinieri Carl, 1917: 392 (D). Thyropygus cuisinieri—Attems, 1930: 157 (D), 1936: 256 (D), 1938: 280 (D, R); Hoffman, 1975: 127 (R); Jeekel, 2006: 14 (R); Enghoff et al., 2004: 36 (R); Enghoff, 2005: 94 (R); Pimvichai et al., 2011b: 41 (D, R). Thyropisthus cuisinieri—Attems, 1942: 79 (D) ; Demange, 1961: 122 (D, R), 1983: 563 (R). Record from Cambodia: Sihanoukville Province, Southeast part of Koh Tang, 10°16.5΄ N, 103°8.9΄E (Pimvichai et al. 2011b). Also known from Vietnam (Carl 1917; Demange 1961; Enghoff et al. 2004; Enghoff 2005; Pimvichai et al. 2011b) and Thailand (Demange 1983; Pimvichai et al. 2011b).

Order Polydesmida

Family Paradoxosomatidae

Genus Helicorthomorpha Attems, 1914

15. Helicorthomorpha luzoniensis (Peters, 1864) Helicorthomorpha luzoniensis—Golovatch, 2015: 128 (R). Record from Cambodia: Kep Province, near Kep (Golovatch 2015). Apparently, a synanthrope species also known from the Philippines, Indonesia (including New Guinea) and China (Golovatch 2015).

16. Orthomorpha coarctata (De Saussure, 1860) Polydesmus coarctatus De Saussure, 1860: 297 (D). Orthomorpha coarctata—Attems, 1953: 179 (R). Record from Cambodia: Koh Kong Province, Sre Ambel (Attems 1953). This is a pantropical anthropochore (e.g. Likhitrakarn et al. 2011).

17. Orthomorpha hydrobiologica Attems, 1930 Orthomorpha hydrobiologica Attems, 1930: 120 (D). Orthomorpha hydrobiologica—Attems, 1937: 63 (D), 1938: 215 (R); Jeekel, 1963: 265 (M), 1964: 361 (M, D), 1968: 45 (M); Hoffman, 1973: 362 (M), 1977: 700 (M); Golovatch, 1998: 42 (M); Enghoff et al., 2004: 38 (R); Likhitrakarn et al., 2011: 53 (D). Oxidus hydrobiologicus—Chamberlin, 1945: 10 (R). Record from Cambodia: Sihanoukville Province, Ream; Kampot Province, Phnom Bokor (Attems 1938). Also known from Indonesia (Attems 1930; Chamberlin 1945) and Vietnam (Attems 1938; Enghoff et al. 2004).

18. Orthomorpha cambodjana (Attems, 1953) Pratinus cambodjanus Attems, 1953: 168 (D). Orthomorpha cambodjana—Jeekel, 1963: 265 (M), 1964: 361 (M, D), 1968: 56 (M); Hoffman, 1977: 700 (M); Golovatch, 1998: 42 (M, D); Likhitrakarn et al., 2011: 66 (D), 2014b: 7 (D, R). Records from Cambodia: Kampot Province, Kampot; Sihanoukville Province, Ream; Koh Kong Province, Sre Ambel (Attems 1953). Also known from Laos (Likhitrakarn et al. 2014b).

Family Cryptodesmidae

Genus Niponia Verhoeff, 1931

19. Niponia kometis (Attems, 1938) Niponielle kometis Attems, 1938: 244 (D). Onomatoplanus kometis—Attems, 1953: 179 (R). Niponia kometis—Enghoff et al., 2004: 41 (R); Likhitrakarn et al., 2014a: 479 (R). Record from Cambodia: Kratié Province, Kratié (Attems 1953). Also known from Laos and Vietnam (Attems 1953; Enghoff et al. 2004).

Conclusions

The millipede fauna of Cambodia currently comprises some 19 species from 15 genera, 12 families and 8 orders. The collecting localities for the millipedes of Cambodia are only very few (Map 1), especially so compared to the neighbouring countries such as Thailand, Laos and Vietnam. Only 3 species are presumably endemic to this country, but the degree of endemism can be expected as eventually being high. For example, Vietnam enjoys at least 90% endemicity at the species level, as opposed to about 80% recorded in Thailand (Likhitrakarn et al. 2014a). The Oriental realm is the globe’s sole biogeographic region that harbours all 16 orders of Diplopoda (Shelley & Golovatch 2011). Thus, based on the available information from the adjacent parts of China, Myanmar, Thailand, Vietnam and/or Laos (e.g. Attems 1938, 1953; Enghoff et al. 2004; Enghoff 2005, Shelley & Golovatch 2011, Likhitrakarn et al. 2014a), the orders Glomerida, Platydesmida, Polyzoniida, Callipodida and Chordeumatida are nearly certain to occur in Cambodia, maybe also Stemmiulida and Siphonocryptida, but none have been recorded yet.

MILLIPEDES OF CAMBODIA Zootaxa 3973 (1) © 2015 Magnolia Press · 181 At the moment we may roughly estimate the Cambodian millipede richness at least at 100 species, i.e. quite comparable to what has hitherto been documented from Thailand (171 species), Vietnam (136 species) or Laos (47 species) (Enghoff et al. 2004; Enghoff 2005; Likhitrakarn et al. 2014a, 2014c, 2014d, 2015).

Acknowledgements

This project was partly funded through grants received from the Office of the Royal Development Projects Board, Office of Agricultural Research and Extension Maejo University, Chulalongkorn University Graduate School Postdoctoral Project to NL, while most of the financial support was received from The Thailand Research Fund, The TRF Senior Research Scholar RTA 5580001 (2012–2015) to SP. We thank the members of the Animal Systematics Research Unit for their invaluable assistance in the field.

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184 · Zootaxa 3973 (1) © 2015 Magnolia Press LIKHITRAKARN ET AL. Zootaxa 4044 (1): 130–140 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2015 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.4044.1.7 http://zoobank.org/urn:lsid:zoobank.org:pub:917B38A3-49BA-4509-AED1-729B78A1EB18 Three new species of the pill millipede genus Hyleoglomeris Verhoeff, 1910, from northern Thailand (Diplopoda, Glomerida, Glomeridae)

NATDANAI LIKHITRAKARN1, SERGEI I. GOLOVATCH2,4& SOMSAK PANHA3,4 1Division of Plant Protection, Faculty of Agricultural Production, Maejo University, Chiang Mai 50290, Thailand 2Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky pr. 33, Moscow 119071, Russia 3Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand 4Corresponding authors. E-mail: Somsak Panha ([email protected]); Sergei I. Golovatch ([email protected])

Abstract

Seven species of the basically warm temperate to tropical Eurasian genus Hyleoglomeris are currently known from Thai- land, including three new ones: H. hongkhraiensis sp. n. and H. aurea sp. n. from Chiang Mai Province, and H. cavicola sp. n. from Sukhothai Province. A new distribution map and a key to all Hyleoglomeris species presently known to occur in Thailand are given.

Key words: millipede, taxonomy, Hyleoglomeris, new species, key, Thailand

Introduction

The millipede genus Hyleoglomeris Verhoeff, 1910, is one of the largest and certainly the most widespread in the entire order Glomerida. At the present, it contains 105 nominate species, ranging from Serbia and Greece in the West to Japan and Taiwan in the East, southern mainland China and the Sunda Archipelago (Sulawesi) in the Southeast (Golovatch et al. 2006, 2012; Liu & Tian 2015). This genus has recently been reviewed (Golovatch et al. 2006, 2012), and its nomenclature finally settled (Golovatch et al. 2011a). Fourteen species of Hyleoglomeris have hitherto been known to occur in Indochina, mostly in Vietnam (6 species) (Golovatch et al. 2013), followed by four species each reported from Laos (Likhitrakarn et al. 2014a) and Thailand (Enghoff 2005). Thus, compared to the fauna of mainland China which supports as many as 30 species (Liu & Tian 2015), the diversity of Indochinese Hyleoglomeris seems to be underestimated.

Material and methods

New material was collected from several provinces of northern Thailand between 2008 and 2015 by the third author and members of the Animal Systematics Research Unit, Chulalongkorn University. Specimens were preserved in 75% ethanol, and morphological investigations were carried out in the laboratory using an Olympus stereomicroscope. Digital images of the specimens were taken in the laboratory and assembled using the “CellD” automontage software of the Olympus Soft Imaging Solution GmbH package. In addition, line drawings of telopods and some other structures were prepared. All holotypes, as well as most of the paratypes, are housed in the Museum of Zoology, Chulalongkorn University (CUMZ), Bangkok, Thailand, some duplicates also being donated to the collections of the Natural History Museum of Denmark, University of Copenhagen, Denmark (ZMUC), the Zoological Museum, State University of Moscow, Russia (ZMUM), and the Naturhistorisches Museum Wien, Austria (NHMW), as indicated in the text. Collecting sites were located by GPS using the WGS84 datum. In the catalogue sections, D stands for the original description or subsequent descriptive notes, whereas R stands for a subsequent record or records.

130 Accepted by W. Shear: 20 Oct. 2015; published: 17 Nov. 2015 Taxonomic part

Family Glomeridae Leach, 1815

Genus Hyleoglomeris Verhoeff, 1910

Hyleoglomeris siamensis Silvestri, 1917

Apiomeris (Hyleoglomeris) siamensis Silvestri, 1917: 116 (D). Hyleoglomeris siamensis—Golovatch, 1983: 116 (D). Hyleoglemeris siamensis—Enghoff, 2005: 88 (R).

Remarks. This species has been described from Raheng (2,000 ft), Meetaw, Thailand (Silvestri 1917). This type locality is Me Taw forest (= Mae Tho), drained by the Me Taw Creek, east of Raheng Town, Tak Province (Barton 1914). Presently known only from Thailand.

Hyleoglomeris albicollis Golovatch, 1983

Hyleoglemeris albicollis Golovatch, 1983: 113 (D). Hyleoglemeris albicollis—Enghoff, 2005: 88 (R).

Remarks. This species has been described from Doi Inthanon National Park (1,700 m a.s.l.), Mae Chaem, Chiang Mai Province, Thailand (Golovatch 1983). Presently known only from Thailand.

Hyleoglomeris cremea Golovatch, 1983

Hyleoglemeris cremea Golovatch, 1983: 114 (D). Hyleoglemeris cremea—Enghoff, 2005: 88 (R).

Remarks. This species has been described from Doi Chiang Dao National Park (1,800 m a.s.l.), Chiang Mai Province, Thailand (Golovatch 1983). Presently known only from Thailand.

Hyleoglomeris montana Golovatch, 1983

Hyleoglemeris montana Golovatch, 1983: 112 (D). Hyleoglemeris montana—Enghoff, 2005: 88 (R).

Remarks. This species has been described from Doi Inthanon National Park, summit (2,500 m a.s.l.), Mae Chaem, Chiang Mai Province, Thailand (Golovatch 1983). Presently known only from Thailand.

Hyleoglomeris hongkhraiensis sp. n. Figs 1 & 2.

Holotype male (CUMZ), Thailand, Chiang Mai Province, Doi Saket District, Huai Hong Khrai Royal Development Study Centre, 445 m a.s.l., 18°52'47"N 99°13'22"E, 02/07/2014, leg. N. Likhitrakarn. Paratypes. 8 males, 9 females (CUMZ), 1 male, 1 female (ZMUC), 1 male, 1 female (ZMUM ƿ2712), 1 male, 1 female (NHMW), same locality, together with holotype; 1 female (CUMZ), same locality, 12/11/2015, leg. N. Likhitrakarn. Name. An adjective to emphasize the type locality. Diagnosis. Differs by the thoracic shield being contrasting yellowish against a dark background, the presence

THREE NEW SPECIES OF HYLEOGLOMERIS FROM NORTHERN THAILAND Zootaxa 4044 (1) © 2015 Magnolia Press · 131 of modest granulations laterally on the telopod prefemur and femur, coupled with a subtrapeziform, large, slightly concave, central lobe of the telopod syncoxite. Description. Length of non-stretched, but unrolled specimens ca 6.0–7.4 mm (males), 6.3–9.3 mm (females), width 2.4–3.4 (males), 2.8–4.5 mm (females). Body mostly dark (Fig. 1A–F). Head and collum blackish to dark brownish. Legs and venter dark brownish to light yellowish. Thoracic shield contrasting yellowish, usually with a narrow dark band caudally and a pair of dark paramedian spots flanking the midline. Mid-dorsal spots on each of terga 3 to 11 usually rather parallel-sided, but quite often V-shaped, subtriangular, often with a distinct, more or less wide and complete, yellow to greyish, axial stripe (Fig. 1A, D & E), this latter usually growing broader and clearer on pygidium. Lateral sides of each of terga 3 to 11 also with a pair of large, sublateral, yellow to marbled yellow-brownish spots normally not reaching translucent caudal and lateral edges (Fig. 1A, C–E). Pygidium with a pair of yellow to greyish parallel stripes always reaching a translucent caudal margin. Labrum sparsely setose (Fig. 2A). Gnathochilarium with 2+2 palps of subequal length. Ocellaria blackish, ocelli 6(7)+1, cornea very convex, translucent. Antennae with four evident apical cones, segment 6 ca 1.4–1.5 times as long as high. Organ of Tömösváry oblong-oval, elongate, ca 1.2–1.4 times as long as broad (Fig. 2A).

FIGURE 1. Hyleoglomeris hongkhraiensis sp. n., (A–C) male holotype, (D–F) male paratype. Habitus in dorsal, ventral, lateral, dorsal, dorsal and ventral views, respectively. Scale bar: 0.2 mm.

132 · Zootaxa 4044 (1) © 2015 Magnolia Press LIKHITRAKARN ET AL. FIGURE 2. Hyleoglomeris hongkhraiensis sp. n., (A–C & F–H) male holotype, (D, E & I) male paratype. A head B thoracic shield, lateral view C–E leg 17 F leg 18 G telopod, front view H & I tip of syncoxital horns J telopod, caudal view. Scale bar: 0.2 mm. Scale bar: A & B drawn not to scale; C–J 0.2 mm.

Collum as usual, with two transverse striae (Fig. 2A). Thoracic shield with a small hyposchism field not projecting caudad behind tergal margin. Striae 7–8, mostly superficial, only lower 3–4 lying above schism, one level to schism, remaining 4 below schism, with 4–5 complete, crossing the dorsum (Fig. 2B). Terga 3 and 4 rather broadly rounded laterally. Following terga in front of pygidium faintly concave medially at caudal edge and with 3 striae starting above lateral edge, sometimes middle stria fading away towards midway. Male pygidium very faintly concave medially at caudal edge. Male legs 17 (Fig. 2C–E) particularly strongly reduced, with a rather low to medium-sized, often irregularly

THREE NEW SPECIES OF HYLEOGLOMERIS FROM NORTHERN THAILAND Zootaxa 4044 (1) © 2015 Magnolia Press · 133 rounded coxal lobe and a 4-segmented telopodite. Male legs 18 (Fig. 2F) less strongly reduced, with a more or less narrowly ogival syncoxital notch and a 4-segmented telopodite; femur with a small, setose, caudomedial tubercle (k) near apex. Telopods (= male legs 19) (Fig. 2G–J) as usual, strongly enlarged and stout, with a subtrapeziform, large, roundly emarginated syncoxital lobe flanked by two setose horns, each of the latter being crowned by a subapical setoid filament (Fig. 2G–I). Prefemur and femur modestly granulate laterally (Fig. 2G & J). Caudomedial femoral process prominent, subapically with a rounded subtriangular lobe on caudal face and modest granulations at base (Fig. 2G & J). Caudomedial process of tibia evident, directed mesad. Tarsus rather moderately sigmoid, narrowly rounded apically, with a strong terminal seta. Remarks. The Huai Hong Khrai Royal Development Study Centre where this new species was found was established under the royal initiative in 1982 in the area of Khun Mae Kuang National Forest Reserve, Chiang Mai Province. It covers approximately 8,500 rai (1,360 hectares). H.M. King Bhumibol realized that the area which used to be abundantly forested and rich in natural resources was deteriorating. His Majesty thus ordered the study centre set up with an objective to conduct research and experimentation using appropriate progressive methods which suited the development needs of the Northern Region, especially the conservation of watersheds, reforestation and agricultural development. Nearly 30 years ago, the mobile plantation area was rehabilitated to Dry Dipterocarp Forest as before.

Hyleoglomeris aurea sp. n. Figs 3A–C & 4.

Holotype male (CUMZ), Thailand, Sukhothai Province, Si Samrong District, Wat Tham Rakhang, near forest, 93 m a.s.l., 17°09'51 "N 99°33'30"E, 22/10/2008, leg. C. Sutcharit and S. Panha. Paratypes. 5 males, 2 females, 11 juveniles (CUMZ), 1 male, 1 female (ZMUC), 1 male, 1 female (ZMUM ƿ2713), 1 male, 1 female (NHMW), same locality, together with holotype. Name. To emphasize a golden yellow body, adjective. Diagnosis. Differs by the peculiar colour pattern which seems to be especially similar to that of H. colorata Golovatch, Geoffroy & VandenSpiegel, 2013, from Vietnam (Golovatch et al. 2013), but the body is considerably larger (ca 9.7–13.0 mm versus 5.0–6.0 mm long), coupled with a 4-segmented male telopodite 17 (Fig. 4A & B), and the higher number of ocelli (7(8)+1 versus 6+1). Description. Length of non-stretched, but unrolled specimens ca 9.7–11.7 mm (males), 10.2–13.0 mm (females), width 5.4–6.4 (males), 5.6–6.8 mm (females). Body bright, pattern annulated (Fig. 3A–C), coloration light yellowish to brown-yellowish with rather narrow, regular, contrasting brown bands across third quarter in caudal parts of terga 2–11, only terga 2 and 12 each with particularly narrow bands at caudal margin. Antennae and distal podomeres rusty brown to blackish. Ocelli blackish. Labrum sparsely setose. Gnathochilarium with 2+2 palps of subequal length. Ocellaria blackish, ocelli 7(8)+1, cornea very concex, translucent. Antennae with four evident apical cones, segment 6 ca 2.7–3.2 times as long as high. Organ of Tömösváry oblong-oval, elongate, ca 1.4–1.6 times as long as broad. Collum as usual, with two transverse striae. Thoracic shield with a small hyposchism field not projecting caudad behind tergal margin. Striae 8–9, mostly superficial, only lower 4–5 lying above schism, one level to schism, remaining 5 below schism, with 7–9 complete, crossing the dorsum. Terga 3 and 4 rather narrowly rounded laterally. Following terga in front of pygidium extremely faintly sinuate medially at caudal edge and with three striae starting above lateral edge, sometimes middle stria fading away towards midway. Male pygidium very faintly concave medially at caudal edge. Male legs 17 (Fig. 4A & B) particularly strongly reduced, with a rather low to medium-sized, often irregularly rounded coxal lobe and a 4-segmented telopodite. Male legs 18 (Fig. 4C) less strongly reduced, with a more or less narrowly ogival syncoxital notch and a 4-segmented telopodite; femur with a small, setose, caudomedial tubercle (k) near apex. Telopods (Fig. 4D–F) as usual, strongly incrassate, with a subtrapeziform, large, roundly emarginated syncoxital lobe flanked by two setose horns, each latter crowned by a subapical setoid filament (Fig. 4F). Prefemur

134 · Zootaxa 4044 (1) © 2015 Magnolia Press LIKHITRAKARN ET AL. and femur micropapillate laterally (Fig. 4D & E). Caudomedial femoral process prominent, subapically with a rounded subtriangular lobe on caudal face, densely setose at base (Fig. 4E). Caudomedial process of tibia rather evident, membranous, densely setose; tibial tubercle absent. Tarsus rather moderately sigmoid, narrowly rounded apically, with a strong terminal seta. Remark. The localities of H. aurea sp. n. and H. colorata (Phong Nha, Vietnam) are separated from each other by ca 700 air-km.

FIGURE 3. Hyleoglomeris aurea sp. n., (A–C) male holotype; Hyleoglomeris cavicola sp. n., (D–F), D male paratype, E & F male holotype, habitus in dorsal, ventral, lateral, dorsal, dorsal and ventral views, respectively. Scale bar: 0.2 mm.

Hyleoglomeris cavicola sp. n. Figs 3D–F & 5.

Holotype male (CUMZ),Thailand, Chiang Mai Province, Chom Thong District, Cave Barijainda, 417 m a.s.l., 18°29'57"N 98°40'19"E, 13/10/2009, leg. N. Likhitrakarn, H. Enghoff, C. Sutcharit and S. Panha.

THREE NEW SPECIES OF HYLEOGLOMERIS FROM NORTHERN THAILAND Zootaxa 4044 (1) © 2015 Magnolia Press · 135 FIGURE 4. Hyleoglomeris aurea sp. n., (A & C–F) male holotype, (B) male paratype. A & B leg 17 C leg 18 D & E telopod, front and caudal views, respectively. F tip of syncoxital lobes. Scale bar: A–E 0.2 mm; F drawn not to scale.

Paratypes. 13 males, 39 females (CUMZ), 2 males, 2 females (ZMUC), 2 males, 2 females (ZMUM ƿ2714), 2 males, 2 females (NHMW), same locality, together with holotype. Name. ‘Cavicola’, a compound Latin noun, derived from ‘caverna’ meaning cave, ending in ‘-cola’ meaning inhabitant. The specific epithet refers to the habitat where this new species occurs. Diagnosis. Differs in the unusual, strongly contrasting colour pattern, in which the head, antennae and collum are brown, while the following terga each with a pair of large, paramedian, dark triangles at their caudal margins and an evident light brown to dark axial stripe. Description. Length of non-stretched, but unrolled specimens ca 7.2–8.9 mm (males), 7.4–11.2 mm (females), width 3.4–4.5 (males), 3.8–5.4 mm (females). Coloration very vivid, colour pattern strongly contrasting (Fig. 3D–F); background light yellowish to pallid. Collum light to dark brown. Thoracic shield with a small, paramedian, dark triangle at caudal margin. Terga 3–11 each with a contrasting pair of large, paramedian, dark triangles at caudal margin and an evident light brown to dark axial stripe. Lateral sides of these terga also with a pair of small, sublateral, dark to marbled light brownish spots beside the triangles (Fig. 3D & E). Pygidium with as pair of large, paramedian, dark, inverted triangles at dorsal margin. Antenna and distal podomeres rusty brown to blackish. Ocelli blackish. Labrum and venter light yellowish to pallid. Labrum sparsely setose. Gnathochilarium with 2+2 palps of subequal length. Ocellaria blackish, ocelli 8(7)+1, cornea very convex, translucent. Antennae with four evident apical cones, segment 6 ca 1.9–2.2 times as long as high. Organ of Tömösváry oblong-oval, elongate, ca 1.3–1.5 times as long as broad.

136 · Zootaxa 4044 (1) © 2015 Magnolia Press LIKHITRAKARN ET AL. FIGURE 5. Hyleoglomeris cavicola sp. n., (A & C–F) male holotype, (B) male paratype. A & B leg 17 C leg 18 D & E telopod, front and caudal views, respectively. F tip of syncoxital horn. Scale bar: A–E 0.2 mm; F drawn not to scale.

Collum as usual, with two transverse striae. Thoracic shield with a small hyposchism field not projecting caudad behind tergal margin. Striae 7–8, mostly superficial, only lower 2–3 lying above schism, one level to schism, remaining 4–5 below schism, with 3–4 complete, crossing the dorsum. Terga 3 and 4 broadly rounded laterally. Following terga in front of pygidium rather clearly concave medially at caudal edge and with two striae starting above lateral edge. Male pygidium clearly concave medially at caudal edge. Male legs 17 (Fig. 5A & B) particularly strongly reduced, with a rather low to medium-sized, often irregularly rounded coxal lobe and a 4-segmented telopodite. Male legs 18 (Fig. 5C) less strongly reduced, with a more or less narrowly ogival syncoxital notch and a 4-segmented telopodite; femur with a small, setose, caudomedial tubercle (k) near apex. Telopods (Fig. 5D–F) as usual, strongly incrassate, with a high, subquadrate, roundly emarginated syncoxital lobe flanked by two setose horns, each latter crowned by a subapical setoid filament (Fig. 5F). Prefemur micropapillate laterally (Fig. 5D & E). Caudomedial femoral process prominent, subapically with a rounded subtriangular lobe on caudal face, modest granulations at base (Fig. 5E & E). Caudomedial process of tibia evident, directed mesad. Tarsus rather moderately sigmoid, narrowly rounded apically, with a strong terminal seta. Remarks. This species was found inside Cave Barijainda and in the surrounding dry dipterocarp forest. This large cave in Doi Inthanon Mountain is located near the Mae Klang River.

THREE NEW SPECIES OF HYLEOGLOMERIS FROM NORTHERN THAILAND Zootaxa 4044 (1) © 2015 Magnolia Press · 137 FIGURE 6. Distribution of Hyleoglomeris species in Thailand (7 species): Open diamond: Hyleoglomeris cremea Golovatch, 1983. Filled circle: H. hongkhraiensis sp. n. Filled square: H. albicollis Golovatch, 1983. Open Circle: H. montana Golovatch, 1983. Filled triangle: H. cavicola sp. n. Open triangle: H. aurea sp. n. Open square: H. siamensis Silvestri, 1917.

Key to the species of Hyleoglomeris currently known to occur in Thailand, chiefly based on male characters:

1. Thoracic shield and pygidium cream-brown, each with a pair of dark spots, not contrasting with cream-brown segments 3–11...... H. siamensis - Thoracic shield and pygidium without a pair of contrasting dark spots ...... 2 2. Background coloration of terga 3–11 dark with lighter pattern/markings ...... 4 - Background coloration of terga 3–11 yellowish with a contrasting dark pattern ...... 3 3. Terga 2–11 with contrasting brown bands in caudal parts of terga 2–11 (Fig. 3A & C)...... H. aurea sp. n. - Terga 3–11 with a contrasting pair of paramedian, dark triangles at caudal margin and a dark axial stripe (Fig. 3D & E)...... H. cavicola sp. n. 4. Collum dark...... 5 - Collum yellowish to pallid ...... 6

138 · Zootaxa 4044 (1) © 2015 Magnolia Press LIKHITRAKARN ET AL. 5. Thoracic shield almost all yellowish to pallid...... H. cremea - Thoracic shield almost dark with a yellowish posterolateral corner...... H. albicollis 6. Thoracic shield almost dark with a thin, yellow-whitish, axial stripe...... H. montana - Thoracic shield almost all yellowish to pallid with a wider axial stripe (Fig. 1A–F)...... H. hongkhraiensis sp. n.

Discussion and conclusion

All seven species of Hyleoglomeris, including the above three new ones, seem to be endemic and confined to northern Thailand alone, even though collecting efforts have covered the whole country since 2006. In addition, Glomerida seem to be encountered in Thailand far less frequently than Sphaerotheriida pill-millipedes. Seasonality may have contributed to that. Thus, nearly each of the numerous species of Tylopus Jeekel, 1968 (Paradoxosomatidae, Polydesmida) found sympatric in the Doi Inthanon (10 species) and Doi Suthep national parks (again 10 species, mostly different) shows its own phenological pattern of occurrence which largely fails to overlap with others (Likhitrakarn et al. 2010, 2014b). It also seems remarkable that Hyleoglomeris spp. are mostly confined to mountainous regions. Only H. aurea sp. n. occurs below 400 m in elevation. This may be accounted for by solely the northern parts of Thailand still supporting larger natural forested areas compared to the central and eastern parts of the country. Similarly, the distributions of Glomerida in Taiwan tend to largely be restricted to montane regions which are still nicely forested (Golovatch et al. 2011b). To summarize, at the present the genus Hyleoglomeris already contains 108 described species, seven of which seem to be endemic to and occur only in northern Thailand. Moreover, Indochina, including north-central and western Thailand, is most likely to yield further new species of Glomerida. This holds especially true of Cambodia where no Glomerida have hitherto been recorded (Likhitrakarn et al. 2015).

Acknowledgements

This project was partly funded through grants received from the Office of the Royal Development Projects Board, while most of the financial support was obtained from The Thailand Research Fund, The TRF Senior Research Scholar RTA 5880002 (2015–2018) to SP. We thank the members of the Animal Systematics Research Unit for their invaluable assistance in the field.

References

Barton, C.S. (1914) A short list of birds from the Raheng District. Natural History Bulletin of the Siam Society, 1 (2), 105–109. Enghoff, H. (2005) The millipedes of Thailand (Diplopoda). Steenstrupia, 29 (1), 87–103. Golovatch, S.I. (1983) On several new Glomeridae (Diplopoda) from Indochina. Annales historico-naturales Musei nationalis hungarici, 75, 107–116. Golovatch, S.I., Geoffroy, J.J. & Mauriès, J.P. (2006) Review of the millipede genus Hyleoglomeris Verhoeff, 1910 (Diplopoda, Glomerida, Glomeridae), with descriptions of new species from caves in Southeast Asia. Zoosystema, 28 (4), 887–915. Golovatch, S.I., Hoffman, R.L. & Chang, H.W. (2011a) Identity of Glomeris bicolor Wood, 1865, and the status of the generic names Hyleoglomeris Verhoeff, 1910 and Rhopalomeris Verhoeff, 1906 (Diplopoda, Glomeridae). Tropical Natural History, 11 (1), 1–8. Golovatch, S.I., Mikhaljova, E.V. & Chang, H.W. (2011b) Pill-millipedes (Glomerida, Diplopoda) in Taiwan. Zootaxa, 2477, 1–20. Golovatch, S.I., Liu, W. & Geoffroy, J.J. (2012) Review of the millipede genus Hyleoglomeris Verhoeff, 1910 in China, with descriptions of new species (Diplopod, Glomerida, Glomeridae). Zootaxa, 3358, 1–27. Golovatch, S.I., Geoffroy, J.J. & VandenSpiegel, D. (2013) On several new species of the millipede family Glomeridae from Vietnam (Diplopoda: Glomerida). Arthropoda Selecta, 22 (3), 201–206. Likhitrakarn, N., Golovatch, S.I., Prateepasen, R. & Panha, S. (2010) Review of the genus Tylopus Jeekel, 1968, with descriptions of the five new species from Thailand (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys, 72, 23–68. http://dx.doi.org/10.3897/zookeys.72.744 Likhitrakarn, N., Golovatch, S.I. & Panha, S. (2014a) A checklist of the millipedes (Diplopoda) of Laos. Zootaxa, 3754 (4), 473–482.

THREE NEW SPECIES OF HYLEOGLOMERIS FROM NORTHERN THAILAND Zootaxa 4044 (1) © 2015 Magnolia Press · 139 http://dx.doi.org/10.11646/zootaxa.3754.4.8 Likhitrakarn, N., Golovatch, S.I. & Panha, S. (2014b) Three new species of the millipede genus Tylopus Jeekel, 1968 from Thailand, with additional notes on the species described by Attems (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys, 435, 63–91. http://dx.doi.org/10.3897/zookeys.435.8286 Likhitrakarn, N., Golovatch, S.I. & Panha, S. (2015) A checklist of the millipedes (Diplopoda) of Cambodia. Zootaxa, 3973 (1), 175–184. http://dx.doi.org/10.11646/zootaxa.3973.1.7 Liu, W. & Tian, M. (2015) A checklist of millipede genus Hyleoglomeris Verhoeff, 1910 in mainland, China with descriptions of seven new species (Diplopoda, Glomerida, Glomeridae). Zootaxa, 4032 (1), 103–116. http://dx.doi.org/10.11646/zootaxa.4032.1.5 Silvestri, F. (1917) Contributions to a knowledge of the oriental Diplopoda Oniscomorpha. I. The family Glomeridae. Records of the Indian Museum, 13 (3.9), 103–151.

140 · Zootaxa 4044 (1) © 2015 Magnolia Press LIKHITRAKARN ET AL. A peer-reviewed open-access journal ZooKeys 472: 27–41 (2015) Review of Kronopolites 27 doi: 10.3897/zookeys.472.9001 RESEARCH ARTICLE http://zookeys.pensoft.net Launched to accelerate biodiversity research

Review of the millipede genus Kronopolites Attems, 1914 (Diplopoda, Polydesmida, Paradoxosomatidae), with the description of a new species from Laos

Natdanai Likhitrakarn1, Sergei I. Golovatch2, Somsak Panha3

1 Division of Plant Protection, Faculty of Agricultural Production, Maejo University, Chiang Mai, 50290, Thailand 2 Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky pr. 33, Moscow 119071, Russia 3 Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chu- lalongkorn University, Bangkok, 10330, Thailand

Corresponding authors: Somsak Panha ([email protected]); Sergei I. Golovatch ([email protected])

Academic editor: R. Mesibov | Received 20 November 2014 | Accepted 15 December 2014 | Published 19 January 2015

http://zoobank.org/D6551D45-E760-4C48-B912-2273F2654AE8

Citation: Likhitrakarn N, Golovatch SI, Panha S (2015) Review of the millipede genus Kronopolites Attems, 1914 (Diplopoda, Polydesmida, Paradoxosomatidae), with the description of a new species from Laos. ZooKeys 472: 27–41. doi: 10.3897/zookeys.472.9001

Abstract The millipede genus Kronopolites currently comprises 11 species, including a new species from northern Laos: K. lunatus sp. n. The generic diagnosis is updated, a key given to all known species, and their distributions are mapped.

Keywords Millipede, Kronopolites, new species, key, distribution, Paradoxosomatidae

Copyright Natdanai Likhitrakarn et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 28 Natdanai Likhitrakarn et al. / ZooKeys 472: 27–41 (2015)

Introduction

The flat-back millipede genus Kronopolites Attems, 1914 is widespread in tropical Asia ranging from the Himalayas of Kashmir, India in the west to Taiwan in the east (Fig. 4). This genus belongs to the mainly Southeast Asian tribe Sulcif- erini in the family Paradoxosomatidae which is one of the largest families in the entire class Diplopoda, dominating the millipede fauna of Indo-Australia (Jeekel 1968, Nguyen and Sierwald 2013). The genus Kronopolites currently contains 10 described species (Golovatch 2013a, 2013b, 2014): K. swinhoei (Pocock, 1895), the type-species which is widespread in central and southeastern China, K. acuminatus Attems, 1937, from northern Vietnam, K. formosanus (Verhoeff, 1939), from northern Taiwan, K. biagrilectus Hoffman, 1963, from Jiangxi Province, China, K. fuscocingulatus Jeekel, 1982, from northern Thailand, K. occidentalis Golovatch, 1983, from the Kashmir Himalaya, India, K. montanus Golovatch, 2009, from northern Vietnam, K. rugosus Golovatch, 2013 and K. davidiani Golovatch, 2014, both from Yunnan Province, China, as well as K. semirugosus Golovatch, 2013 from Sichuan Province, China. The present study treats some new material collected in Laos during several field trips. Prompted by the discovery of a new species, the authors have revised the entire genus Kronopolites adding a new diagnosis and updating both the catalogue and key to species. In addition, its distribution is mapped.

Material and methods

Material was collected in northern Laos in 2014 by SP and members of the Animal Systematics Research Unit, Chulalongkorn University. Specimens were preserved in 75% ethanol, and morphological investigations were carried out in the laboratory using an Olympus stereomicroscope. Scanning electron micrographs (SEM) of gonopods coated with gold were taken using a SEM JEOL JSM–5410 LV microscope. The gonopods were then removed from stubs and returned to alcohol after exami- nation. Digital images of freshly fixed specimens were taken in the laboratory and assembled using the “CellD” automontage software of the Olympus Soft Imaging Solution package. In addition, line drawings of gonopod characters were also prepared. The types are housed in the Museum of Zoology, Chulalongkorn University (CUMZ), Bangkok, Thailand. Collecting sites were located by GPS using the WGS84 datum. In the catalogue sections, D stands for the original description, subsequent descriptive notes or appearance in a key, R for a subsequent record or records, and M for a mere mention. Review of Kronopolites 29

Taxonomic part Family Paradoxosomatidae Daday, 1889 Subfamily Paradoxosomatinae Daday, 1889 Tribe Sulciferini Attems, 1898

Genus Kronopolites Attems, 1914

Kronopolites Attems 1914: 219 (D). Kronopolites – Attems 1929: 272 (D); 1931: 113 (D); 1936: 225 (D); 1937: 49 (D); Verhoeff 1939: 274 (D); Takashima 1950: 38 (M); Takakuwa 1954: 30 (D); Hoffman 1963: 579 (D); 1980: 169 (M); Jeekel 1968: 71 (R); 1971: 225 (M); 1982: 243 (M); 1988: 98 (M); Chen et al. 2006: 252 (M); Golovatch 2009: 121 (D); 2013a: 12 (M); Nguyen and Sierwald 2013: 1287 (M). Kansupus Verhoeff 1934: 17 (D), synonymized by Attems (1936: 233). Kansupus – Jeekel 1971: 225 (M); Hoffman 1980: 169 (M). Parakansupus Verhoeff 1939: 273 (D), synonymized by Hoffman (1963: 579). Parakansupus – Jeekel 1971: 230 (M); Hoffman 1980: 169 (M).

Diagnosis. Body medium-sized to large (ca 23–42 mm long, ca 1.6–6.5 mm wide), with 20 segments. Paraterga from poorly to strongly developed, mostly without lateral incisions. Transverse metatergal sulcus distinct. Sterna usually modified, an acute cone often present near each coxa. Sternal lobe or cone(s) between ♂ coxae 4 present or absent. Pleurosternal carinae usually well-developed. Gonopods rather simple to relatively complex; coxites elongate, subcylindrical, distoventrally sparsely setose, without tubercles; prefemoral (= setose) part of telopodite moderate to relatively large, 1/3–1/2 as long as acropodite; femorite rather slender to stout, slightly curved, enlarged distad, with an evident groove on mesal face and a distinct distolateral sulcus demarcating a postfemoral part; the latter typically carrying a fork consisting of two lateral/ventral processes: usually a smaller basal process b with its tip pointed basad to prefemoral part, and a larger, normally suberect or ventrally curved process a; solenophore strongly developed, slender, slightly longer than or nearly as long as femorite, strongly curved mesad, sometimes with a membranous, distally strongly expanded end, almost completely sheathing a flagelliform and longer solenomere; semi- nal groove running entirely or mostly mesally along an excavate femorite, then directed slightly dorsad in distal part of femorite to follow onto solenomere thereafter. Type species. Strongylosoma swinhoei Pocock, 1895, by original designation. Other species included. K. acuminatus Attems, 1937, K. formosanus (Verhoeff, 1939), K. biagrilectus Hoffman, 1963, K. fuscocingulatus Jeekel, 1982, K. occidentalis Golovatch, 1983, K. montanus Golovatch, 2009, K. rugosus Golovatch, 2013, K. semirugosus Golovatch, 2013, K. davidiani Golovatch, 2014, K. lunatus sp. n. Remarks. Pocock (1895) described the type species in Strongylosoma Brandt, 1833, from a single female from Chee Foo, China. Soon after that Brölemann (1896), 30 Natdanai Likhitrakarn et al. / ZooKeys 472: 27–41 (2015) having received a male of this species from Chou-San Island, China, gave a more de- tailed description, including that of gonopod structure. Attems (1914) proposed a new genus, Kronopolites, and designated Strongylosoma swinhoei as type species.

Kronopolites acuminatus Attems, 1937

Kronopolites acuminatus Attems 1937: 52 (D). Kronopolites acuminatus – Attems 1938: 227 (D); Jeekel 1968: 59 (R); Enghoff et al. 2004: 38 (M, R); Golovatch 2009: 121 (D); Nguyen and Sierwald 2013: 1287 (M). Kronopolites acuminatus acuminatus – Hoffman 1963: 584 (M, R); Golovatch 1983a: 181 (M); Enghoff et al. 2004: 38 (M, R).

Remarks. This species was described from Hagiang, Hagiang Province, Vietnam (At- tems 1937), later redescribed from the type locality (referred to as Ha Giang, 22°50'N, 105°E, 20 miles south of the Vietnam-China frontier) (cf. Hoffman 1963).

Kronopolites biagrilectus Hoffman, 1963

Kronopolites acuminatus biagrilectus Hoffman 1963: 584 (D). Kronopolites acuminatus biagrilectus – Wang and Mauriès 1996: 86 (M); Enghoff et al. 2004: 38 (M); Jeekel 1968: 71 (M); Sierwald 2009: 125 (M). Kronopolites biagrilectus – Golovatch 2009: 121 (D); Nguyen and Sierwald 2013: 1287 (M).

Remarks. This species was described from Kuling, 29°30'N, 116°E, 10 miles south of Kiukiang, Kiangsi (= Guangxi) Province, China (Hoffman 1963).

Kronopolites davidiani Golovatch, 2014

Kronopolites davidiani Golovatch 2014: 10 (D).

Remarks. This species has been described from near Wenchian, 3365 m a.s.l., 27°20'35"N, 99°52'34"E, 214 National Road, Yunnan (not Sichuan!) Province, China (cf. Golovatch 2014).

Kronopolites formosanus (Verhoeff, 1939)

Kronopolites (Parakansupus) formosanus Verhoeff 1939: 273 (D). Kronopolites formosanus – Attems 1940: 540 (D); Takashima 1950: 38 (R); Chamberlin and Wang 1953: 5 (R); Wang 1964: 69 (M); Takakuwa 1954: 31 (D); Hoffman Review of Kronopolites 31

1963: 585 (D); Jeekel 1968: 71 (M); Golovatch 1983b: 298 (M); 2009: 121 (D); Chen et al. 2006: 259 (D); Nguyen and Sierwald 2013: 1287 (M). Kronopolites ralphi Wang 1957: 106 (D), synonymized by Hoffman (1963: 585). Kronopolites ralphi – Wang 1958: 342 (R); 1964: 69 (M).

Remarks. This species had been erroneously listed as a synonym of Kronopolites swinhoei by Wang and Mauriès (1996: 86) and by Korsós (2004: 23) until these mistakes were corrected by Chen et al. (2006). In fact, K. formosanus is endemic to northern Taiwan (Verhoeff 1939, Chamberlin and Wang 1953, Wang 1957), occurring below 1000 m a.s.l.: FuShan Botanical Garden, 726 m a.s.l., Ulai, Taipei County; Yang Ming Shan National Park, ca. 750 m a.s.l., near YuYouRen Tomb, Taipei City, Taiwan (Chen et al. 2006).

Kronopolites fuscocingulatus Jeekel, 1982

Kronopolites fuscocingulatus Jeekel 1982 (D): 238 (D). Kronopolites fuscocingulatus – Enghoff 2005: 97 (R); Golovatch 2009: 121 (D); Nguyen and Sierwald 2013: 1288 (M).

Remarks. Jeekel (1982) described this species from several places in northern Thai- land: Hakka village, 50 km N of Chiang Rai City, 800–900 m a.s.l.; Mac Chan (= Mae Chan), Mae Chan District, Chiang Rai Province; Doi Suthep National Park, Chiang Mai Province. Later, Enghoff (2005) reported new specimens of this species in his checklist: Doi Pha Hom Pok National Park, Northwest of Fang, 1550–1660 m a.s.l.; limestone area, 1300 m a.s.l., Doi Chiang Dao National Park; Kontathan (= Montha Than) Waterfall area, Doi Suthep National Park, Chiang Mai Province.

Kronopolites montanus Golovatch, 2009

Kronopolites montanus Golovatch 2009: 121 (D). Kronopolites montanus – Nguyen and Sierwald 2013: 1288 (M).

Remarks. This species was described from Hoang Lien National Park, ca 2000 m a.s.l., west of Sapa, Lao Cai Province, Vietnam (Golovatch 2009).

Kronopolites occidentalis Golovatch, 1983

Kronopolites occidentalis Golovatch 1983b: 297 (D). Kronopolites occidentalis – Golovatch 1984: 328 (R); 2009: 121 (D); Nguyen and Sier- wald 2013: 1288 (M); Shelley 2014: 3 (R). 32 Natdanai Likhitrakarn et al. / ZooKeys 472: 27–41 (2015)

Remarks. This species was described from Pir Panjal Mountains, 2600 m a.s.l., Tang- marg, Jammu and Kashmir State, India (Golovatch 1983). New specimens were collected near the ruins of Pari Mahal Monastery, 1500 m a.s.l., Srinagar, Jammu and Kashmir State, India (Golovatch 1984).

Kronopolites rugosus Golovatch, 2013

Kronopolites rugosus Golovatch 2013a: 12 (D). Kronopolites rugosus – Nguyen and Sierwald 2013: 1288 (M); Golovatch 2013b: 311 (M).

Remarks. This species has been described from north of Lijiang, 27°01'N, 100°12'E, 2400 m a.s.l., Yunnan Province, China (Golovatch 2013a).

Kronopolites semirugosus Golovatch, 2013

Kronopolites semirugosus Golovatch 2013b: 311 (D).

Remarks. This species was described from NW of Mianning, 2955 m a.s.l., 28°39'13"N, 101°58'34"E, Sichuan Province, China (Golovatch 2013b).

Kronopolites swinhoei (Pocock, 1895)

Stronglosoma Swinhoei Pocock 1895: 354 (D). Kronopolites Swinhoei – Brölemann 1896: 354 (D); Attems 1898: 304 (D); 1914: 219 (R). Kronopolites swinhoei – Attems 1936: 226 (D); 1937: 51 (D); Chamberlin and Wang 1953: 5 (R); Hoffman 1963: 581 (D); Jeekel 1968: 71 (M); Golovatch 1978: 678 (R); 1983b: 298 (M); 2009: 121 (D); 2013a: 2 (R, M); Wang and Mauriès 1996: 86 (M); Geoffroy and Golovatch 2004: 20 (R); Korsós 2004: 23 (R, M); Chen et al. 2006: 252 (M); Nguyen and Sierwald 2013: 1286 (M). Kronopolites swinhoei swinhoei – Attems 1937: 51 (D). Kansupus svenhedini Verhoeff 1934: 17 (D), synonymized by Hoffman (1963: 581). Kronopolites svenhedini – Attems 1936: 233 (R); 1937: 53 (D); Zhang and Li 1978: 12 (R); Wang and Mauriès 1996: 86 (M). Kansupus svenhedini var. dentiger Verhoeff 1934: 19 (D), synonymized by Hoffman (1963: 581). Kronopolites svenhedini dentiger – Attems 1936: 233 (R); 1937: 54 (D).

Remarks. This species is especially widely distributed in mainland China: Chee Foo (Pocock 1895); Chou San Island (Brölemann 1896); Lan Tschou, Gansu (Attems Review of Kronopolites 33

1936); Pei-shui-ho, 700 m a.s.l., northeastern Sichuan and southern Gansu (Attems 1937); Wenchow (= Yung-chia), Chekiang Province; Chekiang Province (Cham- berlin and Wang 1953), Hangchow, Chekiang Province (Hoffman 1963); Taibai Shan Mountains, 1300–1700 m a.s.l.; southern slopes, above Houshenzi, 33°51'N, 107°50'E, Shaanxi Province; Bei Shan National Park, 36°56'N, 102°39'E, ca 90 km NE of Xining, Gansu (not Qinghai) Province (corrected here versus Golovatch 2013a); Grotte du Cirque (Circus Cave), Zheng Xiong County, Yunnan Province; Cave Yan Bao Dong, Zheng Xiong County, Yunnan Province; Cave Ha Chong Dong, near Xingren Huawu, Guizhou Qianxi Province, China (Geoffroy and Golovatch 2004).

Kronopolites lunatus sp. n. http://zoobank.org/BCEF7CDD-7BE6-4B47-B02D-5FA3F658A242 Figs 1–3

Holotype ♂, Laos, Xieng Khouang Province, Phookood District, Cave Pra, ca 1180 m a.s.l., 19°30'02"N, 102°52'20"E, 02.07.2014, leg. R. Srisonchai. Paratype. 1 ♂, Laos, Luang Prabang Province, Chomphet District, Kacham Wa- terfall, ca 440 m a.s.l., 19°38'57"N, 102°04'52"E, 01.07.2014, leg. C. Sutcharit. Name. To emphasize the lateral crescent-shaped processes on the gonopod. Diagnosis. Superficially very similar toK. acuminatus, but differs in the smaller size, the width of midbody pro- and metazonae being 2.4–2.5 and 3.1–3.2 mm, respectively (versus 4.5 mm and 6.5 mm, respectively); tarsal brushes are present until ♂ leg 9 (versus absent), and gonopod process b is > 2 times as long as process a (versus shorter), process a being clearly curved (versus nearly straight) while process b is enlarged and lies adjacent to the femorite (versus clearly separated from the femorite). Eventually, it keys out closest to K. formosanus (see Key below). Description. Length 28.4–29.5 (♂), width of midbody pro- and metazonae 2.4– 2.5 and 3.1–3.2 mm (♂), respectively. Live coloration mostly dark, blackish brown; antennae and head dark brown to light brown, venter and a few basal podomeres light brown to yellow-brown; coloration of alcohol material after four months of preservation faded to dark brown; antennae and epiproct light brown to light yellow, venter and a few basal podomeres light brown to pallid (Fig. 1A–I). Clypeolabral region and vertex densely setose, epicranial suture distinct. Antennae moderately long (Fig. 1A), extending behind body segment 3 (♂) when stretched dorsally. In width, segment 4 < 3 < head < 5 < collum < segment 2 < 6–17 (♂); thereafter body gently and gradually tapering. Collum with three transverse rows of setae: 4+4 anterior, 3+3 intermediate and 4+4 posterior; lateral incisions absent; caudal corner of paraterga very broadly rounded, declined ventrad, produced behind rear tergal margin (Fig. 1A, B). Tegument smooth and shining, prozonae finely shagreened, metaterga finely rugulose (Fig. 1A, C, F); surface below paraterga finely microgranulate (Fig. 1B, D, E). Postcollum metaterga with two transverse rows of setae: 3+3 in anterior (pre- 34 Natdanai Likhitrakarn et al. / ZooKeys 472: 27–41 (2015)

Figure 1. Kronopolites lunatus sp. n., ♂ paratype. A, B anterior part of body, dorsal and lateral views, respectively C segments 10 and 11, dorsal view D segments 9–11, lateral view E–G posterior part of body, lateral, dorsal and ventral views, respectively H, I sternal cones between coxae 4, subcaudal and sublateral views, respectively. sulcus) and 3+3 in posterior (post-sulcus) row, traceable as insertion points. Tergal setae long and slender, mostly abraded, about 1/3 as long as metaterga. Axial line barely traceable both on pro- and metazonae. Paraterga strongly developed (Fig. 1A–F), lying rather high (at upper 1/3 of body), slightly upturned, but lying below dorsum; anterior edge broadly rounded and narrowly bordered, fused to callus; caudal corner very narrowly rounded, starting from segment 15 extending increasingly well beyond rear tergal margin (Fig. 1E, F); lateral edge without incisions (Fig. 1A, C, F); posterior edge nearly straight. Calluses on paraterga narrow, delimited by a sulcus both dorsally and ventrally. Ozopores evident, lateral, lying in an ovoid groove at about 1/4 in front of posterior edge of metaterga. Transverse sulcus usually distinct (Fig. 1A, C, F), slightly incomplete on segment 19, complete on metaterga 3–18 (♂), narrow, line-shaped, shallow, reaching bases of paraterga, faintly ribbed at bottom. Stricture between pro- and metazonae evident, broad and deep, ribbed at bottom down to base of paraterga (Fig. 1A–F). Pleurosternal carinae complete crests with a sharp caudal tooth on segments 2–7, thereafter increasingly strongly reduced until segment 17 (♂). Epiproct (Fig. 1E–G) conical, flattened dorsoventrally, with two small apical papillae; Review of Kronopolites 35

Figure 2. Kronopolites lunatus sp. n., ♂ holotype, right gonopod. A–D mesal, lateral, subcaudal and suboral views, respectively. Scale bars: 0.1 mm. tip subtruncate; pre-apical papillae small, lying close to tip. Hypoproct roundly subtriangular, setiferous knobs at caudal edge small and well-separated (Fig. 1G). Sterna densely setose, without modifications, but with two small, rounded, fully separated, setose cones between ♂ coxae 4 (Fig. 1H, I). Legs rather long and slender, midbody ones ca 1.2–1.3 (♂) as long as body height (Fig. 1A, B, F, G); prefemora without modifications, tarsal brushes present until ♂ leg 9. Gonopods (Figs 2, 3) rather complex; coxa a little curved caudad, sparsely setose distoventrally. Prefemur densely setose, about 1/3 as long as femorite + postfemoral part. Femorite rather stout, with an evident mesal groove and a strong distolateral sulcus demarcating a postfemoral part; the latter well-developed, with very prominent, bipartite, crescent-shape, lateral processes: process a rather short, coiled and pointed; process b long and coiled, also pointed; solenophore clearly curved, long, expanded distomesally, trifid, lamina medialis supporting a long flagelliform solenomere. Remarks. This is the first Kronopolites to be found in Laos. 36 Natdanai Likhitrakarn et al. / ZooKeys 472: 27–41 (2015)

Figure 3. Kronopolites lunatus sp. n., ♂ holotype, right gonopod. A–D right gonopod, mesal, lateral, oral and caudal views, respectively. Scale bar: 0.2 mm.

Key to the species of Kronopolites, chiefly based on♂ characters (modified after Golovatch 2009)

1 Coloration with a contrasting pattern, some parts of body segments being much paler, some other ones much darker...... 2 – Coloration rather uniformly brown to brown-blackish, only venter and legs largely yellowish (Fig. 1A–I)...... 8 2 Paraterga relatively poorly developed, set low (mostly at about upper 1/3 of segments), caudal corners of midbody paraterga usually not projecting behind rear tergal margin, at most narrowly rounded (Fig. 1C, D)...... 3 Review of Kronopolites 37

– Paraterga usually relatively well developed, mostly set higher, caudal corners of midbody paraterga produced behind rear tergal margin, acuminate...... 6 3 Sternal cones on ♂ coxae 4 missing; processes a and b of gonopod nearly independent, slender and long. Northern Thailand...... K. fuscocingulatus – Sternal cones on ♂ coxae 4 present, processes a and b of gonopod on a broad common stem, shorter. China...... 4 4 Surface of metaterga rather smooth; gonopod femorite slender, process a longer, process b shorter, beak-shaped...... K. swinhoei – Surface of metaterga rugose; gonopod femorite stout, processes a and b of gonopod different...... 5 5 Process a of gonopod short and spiniform, process b large and axe-shaped...... K. rugosus – Processes a and b of gonopod subequal in length, ribbon-shaped...... K. semirugosus 6 Coloration dark brown with yellow paraterga; sternal cones between ♂ coxae 4 missing; processes a and b of gonopod short and small, sharing a very distinct common stem; Kashmir Himalayas...... K. occidentalis – Colour pattern different, rear halves of prozonae and fore halves of metazonae usually being black-brown, remaining parts yellowish; sternal cones between ♂ coxae 4 present; processes a and b of gonopod longer and slenderer, their shared base far less conspicuous...... 7 7 Process a of gonopod somewhat shorter than process b. Northern Vietnam...... K. acuminatus – Process a of gonopod somewhat longer than process b. Jiangxi Province, China...... K. biagrilectus 8 Paraterga relatively well developed (Fig. 1A, C, F); pleurosternal carinae evident in ♂ segments 2–16; process a of gonopod clearly shorter than b...... 9 – Paraterga rather poorly developed; pleurosternal carinae evident until ♂ segment 10 at most; processes a and b of gonopod subequal in length...... 10 9 Sternal cones between ♂ coxae 4 present; ♂ tarsal brushes missing; solenophore with conspicuous bipartite, complex, apical processes..... K. montanus – Sternal cones on ♂ coxae 4 missing; ♂ tarsal brushes present until legs of segment 17; solenophore simple and slender, with a little branch set off before apex...... K. davidiani 10 Sternal cone between ♂ coxae 4 single, large. Northern Taiwan....K. formosanus – Two small sternal cones between ♂ coxae 4 (Fig. 1H, I). Northern Laos...... K. lunatus sp. n.

Conclusions

To date, 11 species have formally been described in Kronopolites, mostly found in China (5 species) and northern Vietnam (2 species). Only a single species each has been 38 Natdanai Likhitrakarn et al. / ZooKeys 472: 27–41 (2015)

Figure 4. Distribution of Kronopolites (11 species). Inverted filled triangle, more or less from west to east: K. occidentalis Golovatch, 1983; Filled circle: K. fuscocingulatus Jeekel, 1982; Open circle: K. semirugosus Golovatch, 2013; Asterisk: K. davidiani Golovatch, 2014; Filled square: K. rugosus Golovatch, 2013; Open triangle: K. lunatus sp. n.; Crossed square: K. montanus Golovatch, 2009; Cross circle: K. acuminatus Attems, 1937; Open diamond: K. swinhoei (Pocock, 1895); Filled triangle: K. biagrilectus Hoffman, 1963; Open square: K. formosanus (Verhoeff, 1939). reported from northwestern India, northern Thailand, northern Taiwan and northern Laos (Fig. 4). There is little doubt that many more Kronopolites species are to be found in the future.

Acknowledgements

References

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