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The paradoxosomatid genus Touranella Attems, 1937 is recorded from Laos for the first time, with a new species, Touranella champasaksp. nov., described here. The taxonomy of the genus is discussed, an identification key is provided, and the current distribution of all species is mapped.

The hitherto monospecific genus Pleonoporus is revised based on the syntypes of P. robustus Attems, 1938, housed in Naturhistorisches Museum Wien (NHMW), as well as on specimens of what we interpret as a new species of the genus, Pleonoporus tanzanicus sp. nov., collected in Tanzania and housed in the Museum of Nature – Zoology, Leibnitz Institute for the Analysis of Biodiversity Change (ZMH) for more than a century. Both species are described based on habitus and gonopod structures and illustrated with micrographs, whereas scanning electron microscope images are also provided for the new species. This paper further highlights the importance of natural history collections for taxonomic research and studies on species diversity in general.

Although many new species of the millipede genus Nannaria Chamberlin, 1918 have been known from museum collections for over half a century, a systematic revision has not been undertaken until recently. There are two species groups in the genus: the minor species group and the wilsoni species group. In this study, the wilsoni species group was investigated. Specimens were collected from throughout its distribution in the Appalachian Mountains of the eastern United States and used for a multi-gene molecular phylogeny. The phylogenetic tree recovered Nannaria and the two species groups as monophyletic, with Oenomaea pulchella as its sister group. Seventeen new species were described, bringing the composition of the wilsoni species group to 24 species, more than tripling its known diversity, and increasing the total number of described Nannaria species to 78. The genus now has the greatest number of species in the family Xystodesmidae. Museum holdings of Nannaria were catalogued, and a total of 1,835 records used to produce a distribution map of the species group. Live photographs, illustrations of diagnostic characters, ecological notes, and conservation statuses are given. The wilsoni species group is restricted to the Appalachian region, unlike the widely-distributed minor species group (known throughout eastern North America), and has a distinct gap in its distribution in northeastern Tennessee and adjacent northwestern North Carolina. The wilsoni species group seems to be adapted to mesic microhabitats in middle to high elevation forests in eastern North America. New species are expected to be discovered in the southern Appalachian Mountains. National Science Foundation Advancing Revisionary Taxonomy and Systematics grant [1655635]; Virginia Tech Interfaces of Global Change fellowship; American Museum of Natural History; Virginia Tech Graduate Student Assembly Published version This research was funded by a National Science Foundation Advancing Revisionary Taxonomy and Systematics grant (DEB#1655635). We thank the following permitting agencies for granting scientific collection permits: Georgia State Parks, permit #132016, 142018; Kentucky Department of Parks, permit #1708; Kentucky State Nature Preserves Commission, permit #2017-09-05; North Carolina Department of Natural and Cultural Resources, permit #2018_0234; Tennessee Wildlife Resources Agency permit #3914, 1702; Tennessee Department of Environment and Conservation, permit #2017-008; West Virginia Division of Natural Resources, permit #2016.238; Virginia Department of Wildlife Resources, permit #056958; National Parks Service, permit #GRSM-2016-SCI-2369, GRSM-2019-SCI-2369, SHEN-2016-SCI-0018, BLRI-2014-SCI-0033. Thanks go to Bronwyn Williams and Megan McCuller of the North Carolina Museum of Natural Sciences and Kaloyan Ivanov of the Virginia Museum of Natural History for facilitating loans of millipede specimens. We also thank all the collectors who sent us specimens, particularly Curt Harden, Matt Kasson, and Patricia Wooden. We thank Maddie Hellier for assistance in measuring holotype specimens. We thank Nesrine Akkari, Sergei Golovatch, and one anonymous reviewer for their helpful feedback on previous versions of this manuscript. DAH received support from a Virginia Tech Interfaces of Global Change fellowship, a Theodore Roosevelt Memorial Grant from the American Museum of Natural History, and a Graduate Research Development Program award from the Virginia Tech Graduate Student Assembly.

A new millipede species, Paracortina kyrangsp. nov., is described from a cave in Cao Bang Province, northern Vietnam. The new species is diagnosed by having an extraordinarily long projection on the head of males, reduced eyes, a gonocoxite with two processes, a long and slender gonotelopodite with two long, clavate prefemoroidal processes densely covered with long macrosetae apically, and with a distal, reverse, short spine on mesal side, and a rather sinuous distal part of the telopodite. This is the third species of the genus that is known from Vietnam. A brief comparison of some secondary sexual characters is made.

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: Colchiobrachyiulus montanus Vagalinski, sp. nov., Iraniulus tricornis Vagalinski, sp. nov., Omobrachyiulus armatus 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., Svaniulus ryvkini 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, Colchiobrachyiulus dioscoriadis (Lignau, 1915) and Cyphobrachyiulus redikorzevi (Lohmander, 1936), respectively, listed as comb. nov.Omobrachyiulus brachyurus (Attems, 1899) is formally established as a junior subjective synonym of O. caucasicus (Karsch, 1881), syn. nov., and Omobrachyiulus implicitus ritsensis (Golovatch, 1981) is formally synonymised with the typical Omobrachyiulus implicitus (Lohmander, 1936), syn. nov.Omobrachyiulus sevangensis (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 genus Glyphiulus Gervais, 1847 are described and illustrated. The first species, G. longussp. 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. promdamisp. 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.

A new genus of the millipede family Pseudospirobolellidae, Siliquobolellusgen. nov., is described from limestone mountains in Thailand, based on three new species, viz. Siliquobolellus amicusdraconisgen. et sp. nov. from Uthaithani Province, Siliquobolellus constrictusgen. et sp. nov. from Prachuap Khiri Khan Province and Siliquobolellus prasankokaegen. et sp. nov. (type species) from Lampang Province. The descriptions are based on gonopod morphology and mitochondrial DNA data (COI barcodes). The COI barcodes grouped the three new species in a well-supported Siliquobolellusgen. nov. clade. The mean interspecific COI sequence divergence among the three new species was 12% (range: 8–15%). The mean intergeneric COI sequence divergence between Siliquobolellusgen. nov., Coxobolellus Pimvichai, Enghoff, Panha & Backeljau, 2020, and Pseudospirobolellus Carl, 1912 was 19% (range: 14–23%). Three conspicuous gonopodal synapomorphies differentiate Siliquobolellusgen. nov. from other pseudospirobollellid genera: (1) the telopodital part of the posterior gonopod forms a deep concavity, (2) the telopodite of the anterior gonopod is directed distad and does not reach the tip of the coxal part of the anterior gonopod, and (3) the tip of the anterior gonopod coxa is narrowed, curving mesad. As such, the monophyly of the new genus is well supported by both morphological and mitochondrial DNA data. A distribution map and an identification key to the species are provided.

The millipede genus Cherokia Chamberlin, 1949 is a monospecific taxon, with the type species Cherokia georgiana (Bollman, 1889). The last revision of the genus was made by Hoffman (1960) where he established three subspecies. Here we used molecular phylogenetics to assess the genus and evaluate whether it is a monophyletic group, and if the subspecies are each monophyletic. We included material from literature records and three natural history collections. Newly collected samples were obtained through a citizen science project. Morphological characters underlying subspecies groups???the shape of the paranota, body size, and coloration???were evaluated. A molecular phylogeny of the genus was estimated based on DNA sequences for seven gene loci, and a species delimitation analysis was used to evaluate the status of the subspecies. The documented geographical range of Cherokia in the United States was expanded to include a newly reported state record (Virginia) and about 160 new localities compared to the previously known range. Morphological characters, which included the shape of the paranota and body size that had been historically used to establish subspecies, showed clinal variation with a direct relationship with geographical distribution and elevation, but not with phylogeny. Coloration was highly variable and did not accord with geography or phylogeny. The phylogeny recovered Cherokia as a monophyletic lineage, and the species delimitation test supported the existence of a single species. The subspecies Cherokia georgiana ducilla (Chamberlin, 1939) and Cherokia georgiana latassa Hoffman, 1960 have been synonymized with Cherokia georgiana. The molecular and morphological evidence showed that Cherokia is a monospecific genus with the sole species, Cherokia georgiana, being geographically widespread and highly variable in its morphology. National Science Foundation [1916368] Published version This research was supported by a National Science Foundation grant to P. Marek (Division of Environmental Biology, Systematics and Biodiversity Sciences #1916368). Derek Hennen and Jackson Means helped confirm identifications of Cherokia observations on iNaturalist. We thank those who provided specimens: Roger Birkhead, Sawyer Birkhead, Chris Eaton, Barbara Graham, Christina Fizer, Sherrie White, Jessica Clay, Tracey Muise and Raegan Rainey. Robin Andrews and Bill Shear served on FVV's thesis committee, and provided suggestions for earlier versions of the manuscript. Nesrine Akkari, Weixin Liu and Dragan Antic were reviewers who provided comments on previous versions of this paper.

All cavernicolous species of the millipede family Trichopolydesmidae from the Carpathian-Balkan arch and the Rhodope Mountains have been reviewed. At present the family has been shown to comprise five or six genera with eight or nine species. Two new genera have been described, viz., Balkanodesminusgen. nov., with two new species: B. dentatoidessp. nov. and B. serbicussp. nov., from Bulgaria and Serbia, respectively, and the monospecific Rhodopodesmusgen. nov., with R. niveussp. nov., from Bulgaria. Two new combinations and one new status have been proposed: Balkanodesminus bulgaricus (Strasser, 1962) comb. nov. ex Bacillidesmus bulgaricus Strasser, 1962 and Balkanodesminus dentatus (Strasser, 1966a) comb. nov., stat. nov. ex Bacillidesmus bulgaricus dentatus Strasser, 1966a. All genera and species are diagnosed with the inclusion of the most relevant remarks for each of them. Old museum types are checked for Bacillidesmus filiformis (Latzel, 1884) with lectotype designation, as well as for Trichopolydesmus eremitis Verhoeff, 1898. An identification key to all six genera and a distribution map for the eight species are provided, as well as brief remarks and general considerations on the family Trichopolydesmidae.

A conspicuous Tylopus species is described from Northern Vietnam, namely T. helicorthomorphoides sp. nov. The new species is clearly diagnosed by the gonopodal solenophore completely sheathing the solenomere, both being coiled three times, and the absence of spine z and process h of the gonopod. Fragments of the COI and 16S rRNA genes were extracted, and the phylogenetic analysis also supports the new species.

A new genus of the millipede family Pachybolidae from Southeast Asia is described: Macrurobolusgen. nov., with Spirobolus macrurus Pocock, 1893 as type species. This latter species is DNA barcoded (COI) and redescribed based on male morphological characters, which hitherto were unknown. The new genus differs from other pachybolid genera by having (1) the preanal ring process long and protruding beyond the anal valves and (2) the anterior gonopod telopodite distally abruptly narrowed, forming an extremely long, slender, elevated process curved caudad. Given that Macrurobolusgen. nov. is a monotypic genus, it is aphyletic and thus requires further taxonomic revision.

The Oriental genus Nepalella is reviewed, rediagnosed and shown to comprise 28 species, including N. siamensissp. 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 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.

Male gonopods are useful in taxonomic diagnoses and descriptions of millipedes, although they may vary intraspecifically in shape and size. To assess this intraspecific variation, we used geometric morphometric analysis to compare gonopod morphology among eight isolated populations of the colour-polymorphic southern African millipede Bicoxidens flavicollis. Our results showed that gonopod shapes vary significantly across the examined populations, and elucidated subtle variations. CVA cross-validation test indicates an average classification rate of 75% for the five populations for which we had more than one specimen. Although we had a small number of replicates for three populations, our results still illustrate the importance of applying quantitative approaches to what would otherwise be qualitative and subjective gonopod shape categories in millipedes. As such, the taxonomic assignment of the populations of B. flavicollis may require further investigation, and further revisions would be required with an integrative approach, including molecular data, in order to re-evaluate the taxonomic diversity and distribution data of this species. Finally, we highlight the conservation potential of divergent populations as evolutionary insurance against a dynamic and unpredictable climate, whether or not they undergo full speciation.

Two new millipede species of the genus Coxobolellus Pimvichai, Enghoff, Panha & Backeljau, 2020 from Thailand are described: Coxobolellus saratanisp. nov. from Loei Province and Coxobolellus serratoligulatussp. nov. from Uttaradit Province. The descriptions are based on gonopod morphology and two mitochondrial gene fragments (COI and 16S rRNA). The phylogenetic mtDNA analysis assigned the two new species unequivocally to the consistently well-supported Coxobolellus clade, in which they form a fifth subclade that was well supported by maximum likelihood analysis of 16S rRNA, though neither by Bayesian inference nor by COI. The two new Coxobolellus species share four conspicuous gonopodal synapomorphies of the genus: (1) the protruding process on the coxae of the 3rd (and sometimes 4th) pair of male legs, (2) a large, triangular coxae on the 4th–5th pair of legs, (3) a short process of the preanal ring protruding as far as, or slightly beyond, the anal valves, and (4) the posterior gonopod telopodite divided into two parts, with a conspicuous pore opening at the mesal margin at the end of the coxal part of the posterior gonopod. Thus, the two new species provide further evidence of the well-defined monophyly of the genus Coxobolellus. Finally, the paper provides an updated morphological identification key to all currently described Coxobolellus species.

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. hirtussp. 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. armeniacussp. 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.

This study concerns the diplopod genus Eucampesmella Schubart, 1955, widespread in Brazil. After this work, the genus includes 12 valid species, and three incertae sedis: E. pugiuncula (Schubart, 1946), E. brunnea Kraus, 1959 and E. schubarti Kraus, 1957. The type-species, Eucampesmella tricuspis (Attems, 1931), is redescribed based on the holotype, and the following six new Brazilian species are added: Eucampesmella macunaima sp. nov. from the states of Rondônia, Pará, and Piauí; E. capitu sp. nov. from the states of Piauí and Paraíba; E. brascubas sp. nov. from the state of Sergipe; E. iracema sp. nov. from the state of Pernambuco; E. pedrobala sp. nov. from the state of Ceará; and E. lalla sp. nov. from the state of Rio Grande do Norte. Furthermore, E. lartiguei ferrii (Schubart, 1956) is recognized as a junior synonym of E. lartiguei lartiguei (Silvestri, 1897), which also had its status changed, and E. sulcata (Attems, 1898) is revalidated, prevailing under the name Leptodesmus tuberculiporus Attems, 1898. In addition, drawings, diagnoses, and distribution maps for all species of the genus are provided.