A checklist of the millipedes (Diplopoda) of Cambodia.

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.

Review of the Southeast Asian millipede genus Enghoffosoma Golovatch, 1993 (Diplopoda, Polydesmida, Paradoxosomatidae), with descriptions of new species.

The Southeast Asian millipede genus Enghoffosoma ranges from southern China in the north, western Myanmar in the west, and central Thailand and southern Laos in the south and east. It currently contains seven species, including five new ones, described in this paper: E. zebra sp. n., E. lanceolatum sp. n., E. anchoriforme sp. n., E. funda sp. n. and E. bispinum sp. n. The genus is rediagnosed, a key to all known species is given, and their distributions are mapped.

A checklist of the millipedes (Diplopoda) of Laos.

At the present, the millipede fauna of Laos comprises only 34 species from 20 genera, 13 families and 7 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 Cambodia, the orders Polyxenida, Sphaerotheriida, Chordeumatida, Julida and Polyzoniida must occur in Laos, may be also Glomeridesmida, Siphonocryptida and Stemmiulida, but none has been recorded there yet. Moreover, even some ubiquitous "tramp" species, such as Glyphiulus granulatus (Gervais, 1847), Trigoniulus corallinus (Gervais, 1847), Desmoxytes planata (Pocock, 1895) or Oxidus gracilis (C. L. Koch, 1847), have hitherto not been found in Laos. This shows that a lot more collecting efforts, which have heretofore been rather strongly biased to caves, are required to amass a representative material of Diplopoda of Laos and make it available for study.

The millipede genus Iulidesmus Silvestri, 1895 in Colombia (Polydesmida, Paradoxosomatidae, Catharosomatini).

The paradoxosomatid genus Iulidesmus Silvestri, 1895 in the fauna of Colombia is shown to presently contain nine species. Three of them are still obscure, two of which could be revised based on type (I. alacer (Attems, 1944)) or new material (Orthomorpha proxima Silvestri, 1898), one is a nomen dubium (Strongylosoma glabrum Peters, 1864), while further six are described as new here: Iulidesmus garciae n. sp., I. minerus n. sp., I. florezi n. sp., I. sabaneta n. sp., I. icni n. sp., and I. jeekeli n. sp. A key to all six new species of the genus and their distribution map are provided.

Review of the millipede genus Malayorthomorpha Mrsic, 1996 (Diplopoda, Polydesmida, Paradoxosomatidae), with descriptions of two new species from Thailand and a key to its species.

The millipede genus Malayorthomorpha Mrsic, 1996, so far monospecific and previously known only from Park Belum, Perak State, northern Malaysia, is recorded from a mountain in Betong District, Yala Province, southern Thailand for the first time, being represented there by two new species: M.halabala sp. nov. and M.hulutbeeda sp. nov. Both new species are found to occur syntopically and can be assumed as narrowly endemic to the Titiwangsa Mountain Range which begins in southern Thailand, crosses the Malaysian border, and extends into east and west coast regions of the Malay Peninsula. In addition, the generic diagnosis is slightly updated, and a key to all three species is provided.

The millipede genus Pandirodesmus Silvestri, 1932 in Trinidad and Tobago, Caribbean, with the description of P. jaggernauthi sp. nov. from Trinidad (Diplopoda, Polydesmida, Chelodesmidae, Pandirodesmini).

During several collecting surveys for terrestrial micro-molluscs across Trinidad and Tobago, millipedes of the genus Pandirodesmus were taken by MR on both islands. Samples of P. rutherfordi Shelley Smith, 2015, from Tobago, were recovered in addition to those containing a new species from Trinidad: Pandirodesmus jaggernauthi sp. nov. Additional records, illustrations, and descriptive notes are given for P. rutherfordi. A key is presented to all three species of the genus, and their distributions are mapped.

Two new species of the millipede family Trichopolydesmidae from Bahia state, northeastern Brazil, including a remarkable threatened troglobiont (Diplopoda, Polydesmida).

Phaneromerium troglopterygotum sp. nov. and Moojenodesmus schubarti sp. nov. are described from Bahia state, northeastern Brazil. The former species is a remarkable troglobiont relict endemic to a single limestone cave from the Serra do Ramalho karst area; it shows some clear-cut troglomorphic traits, and particular biology and ecology. The latter species occurs close to limestone outcrops of the Serra do Iuiu karst area. Both regions generally support a typical dry tropical climate, but the cave and outcrop habitats represent sites with high humidity, this being essential for the survival of these and other species.

The Oriental millipede genus Nepalella Shear, 1979, with the description of a new species from Thailand and an updated key (Diplopoda, Chordeumatida, Megalotylidae).

The Oriental genus Nepalella is reviewed, rediagnosed and shown to comprise 28 species, including N.siamensis sp. nov. from southeastern Thailand. All Nepalella species are keyed, and their distributions mapped, being highly localized and mainly allopatric. Unlike most congeners, which are largely confined to subtropical environments (including montane to high-montane conditions, up to 3800 m a.s.l.) or karst caves (eight species, all in southern China alone), the new species is the southernmost in the distribution area of the entire genus, also being among the very few (four) that are restricted to lowland, purely tropical habitats.

Revision of the javanicus species group of the millipede genus Glyphiulus Gervais, 1847, with descriptions of five new species from China (Diplopoda, Spirostreptida, Cambalopsidae).

The javanicus-group of Glyphiulus is re-assessed and its Chinese component species are presently divided between the following two newly-circumscribed species groups, i.e. the formosus- and the sinensis-group. The two can be differentiated, based on the diagnostic characters of the first pair of legs in the male. In addition, metatergal crests being complete and the carinotaxy formula on the collum being I-III+P+M are only characteristic of the formosus-group. A molecular phylogeny of the genus, based on DNA sequencing of four gene fragments of four genes, allows for Glyphiulus to be recovered as a monophyletic group, the phylogenetic relationship being ((Clade A, Clade B), Clade C). Molecular evidence is fully congruent with the morphological one. In addition, based on barcoding data, interspecific p-distances between Glyphiulus species amount to 11.2-24.9%, vs. 0-8.2% for intraspecific p-distances. Five new species of Glyphiulus, all cavernicolous, are described from China: G.sinuatoprocessus Zhao & Liu, sp. nov., G.conuliformis Zhao & Liu, sp. nov. (both from Guangdong Province), G.xiniudong Zhao & Liu, sp. nov., G.scutatus Zhao & Liu, sp. nov. and G.portaliformis Zhao & Liu, sp. nov. (all three from Guangxi Zhuang Autonomous Region). The known Chinese species of the formosus-group appear to mainly be confined to the South China region.

A new genus and two new species of the millipede family Chelodesmidae from Bahia state, northeastern Brazil, including a likely troglobiont (Diplopoda, Polydesmida).

A new genus and two new species of Chelodesmidae are described from two caves of the Joo Rodrigues river system in Bahia state, northeastern Brazil: Strongylosomides troglobius sp. nov. and Rotundotergum elevatum gen. nov., sp. nov. The former species differs from congeners primarily by the gonopods showing the prefemoral process strongly fused caudally along its basal half to the acropodite and, in showing such troglomorphic traits as a nearly unpigmented and fragile body with thin tegument, it seems the first troglobiont to be documented in the genus. Moreover, this new species seems to be so remarkably similar to Leptodesmus stercoriarius Schubart, 1956, from Minas Gerais state, that the following new combination is warranted: Strongylosomides stercoriarius (Schubart, 1956), comb. nov. ex Leptodesmus de Saussure, 1859. The new genus, Rotundotergum gen. nov., seems to be especially similar to Eucampesmella Schubart, 1955, but differs clearly by the absence of a secondary process on the gonopodal prefemoral process, coupled with the latter being unusually long and strongly separated from the acropodite. The type species R. elevatum sp. nov., being partly clearly pigmented, seems to be only troglophilic. The region generally supports a typical dry tropical climate, whereas the caves represent sites with high humidity, this being essential for the survival of these and other species. However, these habitats remain unprotected by Brazilian law, while the Joo Rodrigues River system is threatened by irrigation projects, potential limestone exploitation, asphalt and road widening or construction that pass across the system and can collapse the rock, and pollution of the subterranean drainage. We hope that the new species described here could help us protect their type localities and the Joo Rodrigues River system as a whole.

Identity of the millipede genus Pyrgodesmus Pocock, 1892, the type genus of the family Pyrgodesmidae (Diplopoda, Polydesmida).

Revision of Pyrgodesmus obscurus Pocock, 1892, the only, and type species of Pyrgodesmus Pocock, 1892, shows that the genus currently includes two species, P. obscurus and P. permutatus (Attems, 1936), comb. nov. ex Klimakodesmus Carl, 1932. Both species are properly illustrated and both seem to be endemic to Sri Lanka. Pyrgodesmus is rediagnosed viz--viz the presumably especially similar and likewise oligotypic genus Klimakodesmus from southern India.

Review of the millipede genus Hedinomorpha Verhoeff, 1934, with descriptions of three new species collected in China by Professor Jochen Martens (Diplopoda, Polydesmida, Paradoxosomatidae).

Hedinomorpha martensorum spec. nov., H. jaegeri spec. nov. and H. sympatrica spec. nov. are described, the former species from Qinghai Province, and both latter ones occurring syntopically in Shaanxi Province, China. A brief description and a new diagnosis of Hedinomorpha, a key to its 24 species presently known, and a map showing their distributions are provided. The affinities of Hedinomorpha within the tribe Sulciferini are discussed.

The millipede tribe Brachyiulini in the Caucasus (Diplopoda, Julida, Julidae).

The diplopod tribe Brachyiulini is represented in the fauna of the Caucasus by eight genera and 32 species, of which one genus and 14 species are described as new: Colchiobrachyiulusmontanus Vagalinski, sp. nov., Iraniulustricornis Vagalinski, sp. nov., Omobrachyiulusarmatus Vagalinski, sp. nov., O.fasciatus Vagalinski, sp. nov., O.faxifer Vagalinski, sp. nov., O.kvavadzei Vagalinski, sp. nov., O.lazanyiae Vagalinski, sp. nov., O.ponticus Vagalinski, sp. nov., O.pristis Vagalinski, sp. nov., O.trochiloides Vagalinski, sp. nov., O.unugulis Vagalinski, sp. nov., O.zuevi Vagalinski, sp. nov., Svaniulusryvkini Vagalinski, gen. nov., sp. nov., and S.waltheri Vagalinski, gen. nov., sp. nov.Colchiobrachyiulus Lohmander, 1936, a former subgenus of Megaphyllum, is here elevated to a full genus, and the genus Grusiniulus Lohmander, 1936 is downgraded to a subgenus of the genus Cyphobrachyiulus Verhoeff, 1900, both stat. nov., with their previously described species, Colchiobrachyiulusdioscoriadis (Lignau, 1915) and Cyphobrachyiulusredikorzevi (Lohmander, 1936), respectively, listed as comb. nov. Omobrachyiulusbrachyurus (Attems, 1899) is formally established as a junior subjective synonym of O.caucasicus (Karsch, 1881), syn. nov., and Omobrachyiulusimplicitusritsensis (Golovatch, 1981) is formally synonymised with the typical Omobrachyiulusimplicitus (Lohmander, 1936), syn. nov. Omobrachyiulussevangensis (Lohmander, 1932), originally described in the genus Megaphyllum, is here transferred to the former genus, comb. nov. The diagnoses and descriptions of some genera and subgenera are refined and complemented. A key is given to all genera and species of Brachyiulini that occur in the Caucasus, and their distributions are mapped. Several species are recorded as new to the faunas of Armenia, Azerbaijan, Georgia, or Russia. The distribution patterns of the Caucasian Brachyiulini and their biogeographic implications are discussed.

Two new species of the millipede genus Glyphiulus Gervais, 1847 (Diplopoda, Spirostreptida, Cambalopsidae) from caves in northern Thailand.

Two new species of the genus Glyphiulus Gervais, 1847 are described and illustrated. The first species, G.longus sp. nov., is the second species of the javanicus-group to be found in Thailand. It resembles G.guangnanensis Jiang, Guo, Chen & Xie, 2018, from southern China, but is distinguished by a smaller size and the carinotaxic formula of the collum, combined with ♂ legs 1 bearing very strongly reduced telopodites, the anterior gonopods showing a pair of very long and slender apicomesal processes, and the denser plumose and stout flagella of the posterior gonopods. The second species, G.promdami sp. nov., the fifth member of the granulatus-group in Thailand, seems to be particularly similar to G.subbedosae Likhitrakarn, Golovatch & Panha, 2017, from Laos. However, it can be distinguished from the latter species mainly by showing a uniformly yellow collum and the posterior gonopod coxite bearing several strong setae in median and lateral views, coupled with the anterior gonopod coxosternum being microsetose in the anterior and medial parts in caudal view. An identification key to, and a distribution map of, all seven Glyphiulus species currently known to occur in Thailand are also provided.

Review of the millipede tribe Polydrepanini, with the description of a new species from Kerala state, southern India (Diplopoda, Polydesmida, Paradoxosomatidae, Alogolykinae).

The tribe Polydrepanini is rediagnosed through an elongate, mostly slender, sometimes twisted/helicoid gonofemorite and a more or less thin, mostly flagelliform, suberect, apical or subapical solenomere often forming a conspicuous loop/curve near the base. The tribe presently encompasses seven genera, all redefined, illustrated and keyed: Dasypharkis Attems, 1936, stat. revalid., Delarthrum Attems, 1936, Grammorhabdus Carl, 1932, stat. revalid., Pocockina Jeekel, 1965, Polydrepanum Carl, 1932 (= Gyrodrepanum Carl, 1932 and Hindomorpha Golovatch, 1984, both syn. nov.), Telodrepanum Carl, 1932, and Xiphidiogonus Carl, 1932. The genus Polydrepanum is shown to currently comprise only four described species, all also redefined, depicted and keyed: Polydrepanum tamilum Carl, 1932 (the type species), P. horridum Golovatch, 1984, P. granuliferum (Attems, 1936), comb. nov. ex Hindomorpha, and P. lamprum (Chamberlin, 1920), comb. nov. ex Gyrodrepanum. Grammorhabdus fissus (Sankaran Sebastian, 2018) is formally transferred from Polydrepanum, comb. nov. Because Polydrepanum implicatum Carl, 1941 shows a strongly shortened gonopodal femorite, coupled with a distal and flagelliform solenomere, this species is considered as not only somewhat intermediate between the tribes Alogolykini and Polydrepanini as redefined here, but it is formally assigned to Alogolykini as the type species of Carlogonopus gen. nov. (masculine), with C. implicatus (Carl, 1941), comb. nov. ex Polydrepanum, as well as a comb. nov. ex Telodrepanum. A somewhat similar situation concerns Manikidesmus suriensis Bhakat, 2021, a still invalid genus and species from West Bengal, eastern India that likewise shows a clearly shortened gonofemorite. However, this being even more important, Manikidesmus suriensis is a nomen nudum not available yet for zoological nomenclature. The large and mainly Himalayan genus Delarthrum is briefly reassessed, mainly in connection with a new species, D. anomalans sp. nov., described from Kerala, southern India and placed in a species-group of its own. Some general considerations concerning the diversity and distribution of the subfamily Alogolykinae are presented.

The millipede genera Amblyiulus Silvestri, 1896 and Syrioiulus Verhoeff, 1914 in the Caucasus, with notes on their distributions (Diplopoda, Julida, Julidae).

In the Caucasus, the genera Amblyiulus Silvestri, 1896 and Syrioiulus Verhoeff, 1914 are shown to include two and four species, respectively: Amblyiulus georgicus Lohmander, 1932, from Georgia and Armenia, A. hirtus sp. nov., from Azerbaijan and Dagestan, Russia, Syrioiulus adsharicus (Lohmander, 1936), from Georgia, S. continentalis (Attems, 1903), from Azerbaijan and Iran, S. taliscius (Attems, 1927), from Azerbaijan, and S. armeniacus sp. nov., from Armenia. All these six species are described, illustrated, and keyed, and their distributions are mapped and discussed, based on the literature data and abundant new samples.

Two new species of the millipede genus Tylopus Jeekel, 1968 from Shan State, Myanmar (Diplopoda, Polydesmida, Paradoxosomatidae).

The predominantly Indochinese to southern Chinese millipede genus Tylopus presently comprises 76 described species, including two new, T. monticola sp. nov. and T. sutchariti sp. nov., both described and illustrated based on material from a limestone mountain in Taunggyi District, Shan State, Myanmar. Both new species have been found to occur syntopically near limestone caves and are assumed to be narrowly endemic to the Taunggyi Mountains, southwestern Shan State, Myanmar. A key to all six Tylopus species known to occur in Myanmar is provided, and their distributions are also mapped.

The millipede genus Klimakodesmus Carl, 1932, with the description of a new species from Kerala state, southern India (Diplopoda, Polydesmida, Pyrgodesmidae).

Klimakodesmus Carl, 1932 is briefly redescribed, rediagnosed, and shown to be an oligotypic genus endemic to southern India and distinct from the particularly similar genus Pyrgodesmus Pocock, 1892, monobasic and endemic to Sri Lanka, by several important features of peripheral and, especially, gonopodal structure. A new species, Klimakodesmus bilobocaudatus sp. nov., is described from Kerala state, India, differing from the sole accepted, and type species K. gravelyi Carl, 1932, from Tamil Nadu state, primarily by the laterally trilobate paraterga, the caudally more deeply bilobate mid-dorsal keel on ring 19, and certain minor details of the gonopodal structure.

Diversity, distribution patterns, and fauno-genesis of the millipedes (Diplopoda) of mainland China.

Based on all available information, 339 species from 71 genera, 26 families, and eleven orders of Diplopoda have hitherto been recorded from mainland China, the fauna thus being very rich, albeit far from completely known, comprising various zoogeographic elements and populating very different environments. Diplopods mainly occur in various woodlands, in caves, and high in the mountains. Most species (> 90 %, usually highly localised, including 160 cavernicoles), 18 genera, and one family are strictly endemic to continental China. Mapping not only the horizontal, but also the vertical distributions of Diplopoda in China shows the bulk of the fauna to be expectedly restricted to forested lowland and mountain biomes or their remnants. Yet some Chordeumatida, Callipodida, Polydesmida, Julida, and even Spirobolida seem to occur only in the subalpine to alpine environments and thus may provisionally be considered as truly high-montane. The long-acknowledged notions of China being a great biogeographic zone transitional between the Palaearctic and Oriental regions generally find good support in millipede distributions, in particular at the higher taxonomic levels (generic, familial, and ordinal). While the Palaearctic/Holarctic components expectedly dominate the fauna of the northern parts of the country, the Oriental ones prevail in its south and along the Pacific coast. Both realms are increasingly mixed and intermingled towards China's centre. However, in addition to the above traditional views, based on distribution patterns alone, southern China seems to harbour a rather small, but highly peculiar faunal nucleus or origin centre of its own, whence Himalaya, Myanmar, Thailand, Indochina and/or Taiwan could have become populated by younger lineages. The millipede fauna of continental China is thus a tangled mixture of zoogeographic elements of various origins and ages, both relict and more advanced. The few anthropochores must have been the latest faunal "layer" to populate China.

Some new or poorly-known Zephroniidae (Diplopoda, Sphaerotheriida) from Vietnam.

Three new species of the giant pill-millipede family Zephroniidae are described from southern Vietnam: Sphaerobelum pumatense sp. nov., Sphaeropoeus honbaensis sp. nov. and Sphaeropoeus bidoupensis sp. nov. Two species, Sphaerobelum bicorne Attems, 1938 and Sphaeropoeus maculatus (Verhoeff, 1924), are redescribed, the former from new material, the latter from type material with lectotype designation. A new transfer is proposed: Zephronia manca Attems, 1936, to the genus Sphaeropoeus Brandt, 1833, giving the new combination, Sphaeropoeus manca (Attems, 1936) comb. nov.