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.

Integrative taxonomy reveals two new giant pill-millipedes of the genus Zephronia Gray, 1832 from eastern Thailand (Diplopoda, Sphaerotheriida, Zephroniidae).

A large amount of material of the millipede genus Zephronia Gray, 1832 was collected during 2014-2023 from many parts of eastern Thailand. An integrative study of morphological characters and genetic data (COI gene) revealed two new species: Z.chantaburiensis Srisonchai & Wesener, sp. nov. and Z.macula Srisonchai & Wesener, sp. nov. The two new species clearly differ from other congeners by their unique characteristics, especially in their colour pattern and telopod shape. The interspecific genetic distances of the 658 bp COI gene barcoding fragment between these new species and all other species of giant pill-millipede from Thailand, Laos and Cambodia are 12.01-23.49% for Z.chantaburiensis sp. nov. and 17.93-25.13% for Z.macula sp. nov. While relationships among species remain preliminary, the phylogenetic tree shows that species of Zephronia are interspersed with species of Sphaerobelum Verhoeff, 1924 and Prionobelum Verhoeff, 1924. Phylogenetic analyses place both new species in a clade termed Zephronia s.s., which receives support also from morphological data, showing a unique position of the organ of Tömösváry. Z.macula sp. nov. appears to occur over a broad distribution whereas Z.chantaburiensis sp. nov. was found only at the type locality. Given that all known records are in the eastern part of Thailand, we thus regard both species as endemic. Morphological illustrations based on SEM micrographs and a distribution map are also provided.

First records of penicillate millipedes (Diplopoda, Polyxenidae) from Thailand, with descriptions of two new species.

Two new species are described from the family Polyxenidae. Monographis panhai sp. nov. and Unixenus thailandicus sp. nov. represent the first records of their respective genera in Thailand. Monographis panhai sp. nov. shows distinct morphological differences in the arrangement of sensilla in the antennomere VI, as well as in the structures of the labrum and the claw. Collections from different areas of Thailand showed that not only is it widespread but also exhibits geographic variation in morphology. Unixenus thailandicus sp. nov. exhibits sexual dimorphism in the number of conical sensilla on the lateral palp, with 24 in males and 22 in females, differing from the typical count of 13 within the genus.

A new jewel-like species of the pill-millipede genus Sphaerobelum Verhoeff, 1924 (Diplopoda, Sphaerotheriida, Zephroniidae) from Thailand.

A new species of the giant pill millipede genus Sphaerobelum is described: Sphaerobelumturcosasp. nov. from the northeastern part of Thailand. Species delimitation is based on morphological characters and COI sequence data. The new species can be clearly discriminated from congeners by its greenish-blue body color, the face mask-like appearance of the thoracic and anal shields jointly when rolled up, and the combination of the following four characters: (1) the coxa of the second leg laterally with a sharp and long process, (2) the tarsi of legs 4-21 with 6-7 ventral spines, (3) the anterior telopods consisting of four conspicuous telopoditomeres, and (4) the immovable, slender (not strongly humped) and distally curved finger of the posterior telopods without a membranous spot. The interspecific COI sequence divergence between the new species and other Sphaerobelum species ranges from 17% to 23% (mean 20%). The intergeneric COI sequence divergence between the new species and Zephronia species ranges from 18% to 21% (mean 20%). The relationships among Sphaerobelum and Zephronia species based on the COI sequence data were not resolved in this study. Sphaerobelumturcosasp. nov. is restricted to limestone habitat in Loei province and is probably endemic for the Thai fauna.

Review of the pill millipede genus Hyperglomeris Silvestri, 1917 (Diplopoda, Glomerida, Glomeridae) with description of two new species from Laos.

The pill millipede genus Hyperglomeris Silvestri, 1917 is reported from Laos for the first time. Two new species, namely H.bicaudata Likhitrakarn, sp. nov. and H.inkhavilayi Likhitrakarn, sp. nov., from Houaphanh and Khammouane provinces, northern Laos, are described and illustrated based on morphological characters and molecular analyses. Sequences of COI gene were used as DNA barcoding markers, and successfully supported the accurate identification of other Glomeridae species. Interspecific divergence of the COI uncorrected p-distance between these new species and other Hyperglomeris species ranged from 7.84-13.07%, while the intraspecific divergence was 0.45% in H.inkhavilayisp. nov. and 5.3% in H.bicaudatasp. nov. The updated status of Hyperglomeris, a map of its distribution, and identification keys for all species are given.

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 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.

The giant pill-millipede genus Zephronia Gray, 1832 from Thailand, with a redescription of Z.siamensis Hirst, 1907 and descriptions of three new species (Diplopoda, Sphaerotheriida, Zephroniidae).

Material of the giant pill-millipede genus Zephronia Gray, 1832 recently collected from Thailand contains three new species: Zephroniaenghoffi sp. nov., Zephroniagolovatchi sp. nov., and Zephroniapanhai sp. nov. The first Zephronia species recorded for Thailand, Z.siamensis Hirst, 1907, is also redescribed based on new specimens collected both from the type locality in Chonburi Province and from neighboring areas. Morphological characters of all new species, Z.phrain Likhitrakarn & Golovatch, 2021, and Z.siamensis are illustrated, and a distribution map of the confirmed Zephronia species occurring in Thailand is also provided.

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.

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.

Biodiversity and Spatiotemporal Variation of Longhorn Beetles (Coleoptera: Cerambycidae) in Tropical Forest of Thailand.

Longhorn beetles are highly diversified and important for agriculture and health of the environment. However, the fauna and ecology of these beetles are not well known in Thailand. This study is the first to report the biodiversity, elevation, and seasonal distribution of longhorn beetles. Specimens were collected by malaise traps from 41 localities in 24 national parks throughout the country during 2006-2009. The traps were operated at each site for 12 consecutive months with a monthly service. A total of 199 morphotaxa in 36 tribes of 6 subfamilies were identified from 1376 specimens. Of these, 40.7% and 14.5% of total taxa were singletons and doubletons, respectively. The Shannon diversity index and observed species richness at Panernthung, Loei Forest Unit and Mae Fang Hotspring were high at 0.96 (30), 0.88 (50), and 0.86 (34), respectively. Local richness ranged between 3 and 50 species, while the species richness estimator showed between 6 and 1275 species. The most relatively abundant species, Nupserha lenita, Pterolophia sp.1, Oberea sp.3, Acalolepta pseudospeciosa, and Ac. rustricatrix represented 4.80%, 4.80%, 4.80%, 4.5%, and 4.43% of the species, respectively. The species with the widest distribution range of percentage of species occurrence (% SO) was Pt. sp.1 (63.4%), followed by Ac. rustricatrix (39%) and Moechotypa suffusa (39%). In a significantly negative relationship between species richness and elevation (p > 0.05, R2 = 0.04), the species richness pattern showed a hump-shaped curve that peaked at the middle elevation (501-1000 m asl). Regarding seasonal variation, most of the species occurred during the hot season (March-April) and peaked in early rainy season (May), while a low number of species were found during the mid-rainy (June-October) and cold season (November-February). Ordination analysis indicated that the distribution of most species was associated with regions and forest type, and most of the species correlated with forest located at middle and low elevation. The results of this study indicated the very high biodiversity of longhorn beetles in Thailand, which suggests that an understanding of their seasonal and elevational distribution will be of value to agriculture management and conservation. They also indicated that malaise traps are appropriate for the evaluation of biodiversity.

First record of non-flying mammalian contributors to pollination in a tropical montane forest in Asia.

This study aims to identify the flower visitors of Mucuna thailandica (Fabaceae), endemic plant species in montane forests in Thailand, to determine their potential pollinators. The genus Mucuna produces papilionaceous flowers and has an explosive flower-opening step. Explosive opening rapidly exposes stamens and pistil from keel petals and releases pollen. The flower of this species depends completely on animals to perform this step, essential for pollination success. Using a camera trap survey, we revealed that non-flying mammals, such as squirrels (Callosciurus sp.) and masked palm civets (Paguma larvata), opened flowers explosively. Thus, these mammals contribute to the pollination of M. thailandica. This is the first report of non-flying mammals contributing to pollination in montane forests in tropical Asia.

A remarkable new species of the millipede genus Trachyjulus Peters, 1864 (Diplopoda, Spirostreptida, Cambalopsidae) from Thailand, based both on morphological and molecular evidence.

A new, giant species of Trachyjulus from a cave in southern Thailand is described, illustrated, and compared to morphologically closely related taxa. This new species, T. magnus sp. nov., is much larger than all other congeners and looks especially similar to the grossly sympatric T. unciger Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2012, which is widespread in southern Thailand. Phylogenetic trees, both rooted and unrooted, based on a concatenated dataset of the COI and 28S genes of nine species of Cambalopsidae (Trachyjulus, Glyphiulus, and Plusioglyphiulus), strongly support the monophyly of Trachyjulus and a clear-cut divergence between T. magnus sp. nov. and T. unciger in revealing very high average p-distances of the COI gene (20.80-23.62%).

Two new species of the millipede genus Plusioglyphiulus Silvestri, 1923 from Cambodia (Diplopoda, Spirostreptida).

Two new species of Plusioglyphiulus are described from southern Cambodia. Plusioglyphiulus biserratus sp. nov. is clearly distinguished from all congeners by the shape of the telopodites of the posterior gonopods which are distinctly serrate laterally and by the anterior gonopods showing only a pair of single, smooth and curved coxosternal processes. Plusioglyphiulus khmer sp. nov. is distinguished by having most crests on the collum being complete and male legs 1 showing long, prominent, one-segmented telopodites, coupled with the oblong-subtrapeziform, membranous, posterior gonopods with a small bifid process at about a third of the telopodite length. Notes on the variation of Plusioglyphiulus boutini Mauriès, 1970 are also given, including a colour photograph of fresh, live material. A key to all four species of Plusioglyphiulus currently known to occur in Cambodia is also presented.

A new micropolydesmoid millipede of the genus Eutrichodesmus Silvestri, 1910 from Cambodia, with a key to species in mainland Southeast Asia (Diplopoda, Polydesmida, Haplodesmidae).

The micropolydesmoid millipede family Haplodesmidae is here recorded from Cambodia for the first time through the discovery of the first, new species of the genus Eutrichodesmus Silvestri, 1910: E. cambodiensis sp. nov. This new species is described from two limestone habitats in Kampot Province, based on abundant material. It is easily distinguished from all related congeners by the following combination of characters: body greyish-brown; limbus roundly lobulate; solenomere partially divided from acropodite by a digitiform lobe, but without hairpad. Brief remarks on the previously-proposed "pecularis-group" are provided and a second group, the "demangei-group", is established and discussed on the basis of morphological evidence, updating the number of recognised species groups of Eutrichodesmus to two. Detailed morphological illustrations, photographs and a distribution map, as well as remarks on its habitat and mating behaviour of the new species are presented. Furthermore, the current distributions of all 55 presently-known species of Eutrichodesmus are provided and a key to all 23 species that occur in mainland Southeast Asia is given.

A new species and a new record of the Southeast Asian millipede genus Antheromorpha Jeekel, 1968 (Polydesmida, Paradoxosomatidae) from Vietnam.

Antheromorphanguyeni sp. n. is described and illustrated from Kon Ka Kinh National Park, southern Vietnam. The new species is distinguished by a peculiar colour pattern showing a uniformly black-brown body contrasting with yellow-brown paraterga and epiproct, as well as in the pointed gonopodal process being unusually short, only approximately half as long as the solenophore. In addition, an identification key to all 13 presently known species, all mapped, is given. A new record of A.festiva is provided from southern Vietnam.

A new species of the millipede genus Cryptocorypha Attems, 1907, from northern Thailand (Polydesmida, Pyrgodesmidae).

The millipede family Pyrgodesmidae and the genus Cryptocorypha are recorded from Thailand for the first time, being represented there by C.enghoffi sp. n. The new species is distinguished by the evident apicodorsal trichostele on the last tibia of both sexes and the gonopodal telopodite being particularly complex, quadripartite, consisting of the longest, mesal, suberect solenomere branch; a slightly shorter, similarly slender, acuminate endomere branch tightly appressed to the solenomere; a somewhat shorter, caudal, strongly curved, armed exomere process; and a very distinct, low, lateral, sac-shaped velum at their base. This situation strongly resembles the one observed in the geographically closest C.perplexa Golovatch & VandenSpiegel, 2015, from Myanmar, but the shapes and armament of all outgrowths of the gonopodal telopodite are clearly different. A key to all three Cryptocorypha pecies known from Indochina or Myanmar and an updated checklist of all 21 species of the genus are provided.

Review of the millipede genus Orthomorpha Bollman, 1893 (Diplopoda, Polydesmida, Paradoxosomatidae) in Vietnam, with several new records and descriptions of two new species.

The genus Orthomorpha is shown to currently be represented in Vietnam by ten species or varieties, including new records of O. arboricola (Attems, 1937), O. coarctata (de Saussure, 1860), O. rotundicollis (Attems, 1937) and O. scabra Jeekel, 1964, and two new species, O. caramel sp. nov. and O. vietnamica sp. nov. A key to all eight Orthomorpha species and two varieties known to occur in Vietnam is provided. Although the morphological characters that have been traditionally used for Orthomorpha taxonomy are here considered superior to molecular ones, molecular-based phylogenetic relationships and taxon assignments within the tribe Orthomorphini are provisionally analyzed using fragments of the cytochrome c oxidase subunit I (COI) mitochondrial gene. The preferred phylograms, both rooted and unrooted, demonstrate the monophyly of the tribe Orthomorphini, but due to the special, uncertain or even controversial position of O. coarctata, which occurs closer to the genera Antheromorpha and Hylomus, the genus Orthomorpha in current usage appears to be polyphyletic. However, if O. coarctata is to be treated within the monotypic genus Asiomorpha, the monophyly of Orthomorpha becomes manifest. On the other hand, a cautious approach is followed to avoid descriptions of suspicious new taxa/species. Thus, solely because the average genetic distance between O. rodundicollis subrotundicollis var. nov. and O. rodundicollis, as well as that between O. scabra grandis var. nov. and O. scabra, are both found to be negligibly small, the statuses of the sympatric and closest yet morphologically different varieties are treated only as such, i.e., infrasubspecific categories. The apparent discord observed between morphological and molecular data is obviously due to only partial and single-gene topologies used, possibly also to hybridization already known to occur in some closely related and sympatric paradoxosomatid species or even genera.