The Giant Pill-Millipedes of Nepal (Diplopoda, Sphaerotheriida, Zephroniidae).

The type of the only species of the order Sphaerotheriida with a record in Nepal, Kophosphaera excavata (Butler, 1874), originally described from Sikkim, is redescribed. The subspecies K. excavata mammifera Attems, 1936 from Sureil, Darjeeling, India, is elevated to species rank, K. mammifera stat. nov.. A species of unclear country of origin ('Himalaya'), Sphaeropoeus montanus Karsch, 1881, is briefly redescribed and transferred to the genus Zephronia, Z. montana (Karsch, 1881) n. comb.. Z. tumida Butler, 1882, an apparently widespread north Indian Zephronia species, is redescribed. Sphaerotheriida specimens collected during several expeditions to Nepal undertaken by Prof. J. Martens in the 1970s and 1980s were examined. The material contained 10 specimens (7 males, 3 females) from seven localities, including three undescribed species, Zephronia nepalensis n. sp., Kophosphaera shivapuri n. sp., and Kophosphaera martensi n. sp., as well as a specimen of Kophosphaera politissima Attems, 1935, type species of the genus and described previously from India. A key to all (now seven) species of Kophosphaera is presented. A brief diagnosis of the Kophosphaera excavata group is provided. While Zephronia seems to be restricted to the eastern part of Nepal, two endemic and two more widespread Kophosphaera species occur also in its central and mid-western part, representing the western-most records of the family Zephroniidae in Asia. The current distribution of the family in Nepal clearly indicates the Zephroniidae as a family adapted to tropical environments.

Redescription of 'Polyzonium' malagassum, a new synonym of Rhinotus purpureus (Pocock, 1894), with notes about the occurrence of the order Polyzoniida on Madagascar (Diplopoda).

Polyzonium malagassum de Saussure & Zehntner, 1902, the only indigenous record of the order Polyzoniida from Madagascar, is redescribed after a study of the type specimens. The only male specimen is selected as the lectotype and illustrated. P. malagassum is discovered to be a synonym of the widespread tropical tramp species Rhinotus purpureus (Pocock, 1894). A mapping of additional locality data of R. pupureus shows that the species is widespread in Malagasy rainforests and montane rainforests, and occurs locally in high densities. Seven potentially indigenous Polyzoniida morphospecies also occur on Madagascar, but these undescribed species are more localized and show a lower abundance than R. purpureus. Brief notes, locality data, and Museum acronyms are given for the undescribed Polyzoniida species, which will hopefully assist future studies on Malagasy representatives of this little-known but biogeographically interesting order. With the discovery of the ubiquitous presence of R. purpureus on Madagascar, the similarity of the defense secretions of South American and of endemic Malagasy poison dart frogs (family Mantellidae) might derive from the fact that both groups prey on and sequester alkaloids from the same species of millipede. 

The first record of the giant pill-millipede genus Prionobelum Verhoeff, 1924 from Thailand, with the integrative description of two species (Diplopoda, Sphaerotheriida, Zephroniidae).

Thailand hosts a very rich but underexplored giant pill-millipede (Sphaerotheriida) fauna, with 11 of its 13 species described in the last three years. Currently, all known Thai giant pill-millipedes belong to the genera Zephronia Gray, 1832 (nine species) and Sphaerobelum Verhoeff, 1924 (four species). Here we describe the first two species of the genus Prionobelum Verhoeff, 1924 (previously restricted to Vietnam and China), Prionobelum inthanonense n. sp. and P. naevium n. sp. from Thailand. The species occur at Thailands highest mountain (2500 m) Doi Inthanon and the lowland rainforests at Bang Lang National Park touching the border with Malaysia. Both species are described integratively, utilizing light microscopy, scanning electron microscopy as well as DNA barcoding. Both new species of Prionobelum differ from other Zephroniidae species, as well as from one another, by more than 20% p-distance in the COI barcoding gene suggesting that potential closer related species are still awaiting discovery.

Madagascarhinus, a new genus of the family Siphonorhinidae with two new species from Madagascar (Diplopoda, Siphonophorida).

Millipedes of the family Siphonorhinidae Cook, 1895, famous for including the leggiest animal on this planet until recently, with up to 750 legs, show a disjunct distribution in California, South Africa and South-East Asia (from India to the Indonesian Islands of Java and Flores). Here I formally describe the first members of the family from Madagascar, Madagascarhinus madagascariensis n. gen., n. sp. and Madagascarhinus andasibensis n. gen., n. sp., from degraded central highland rainforests. Both species of Madagascarhinus n. gen. are relatively short (61 or 63 tergites), setose, pale and slender, and differ from the other known five genera of the family in numerous characters of the antennae, head, and tergites.

Integrative redescription of the sucking millipede genus Dawydoffia Attems, 1953, with a description of a new species and a transfer to the family Hirudisomatidae (Diplopoda, Polyzoniida).

The type species of the monotypic Polyzoniida genus Dawydoffia Attems, 1953, D. kalonota Attems, 1953 from Vietnam, is redescribed based on type material. A second species of the genus, D. siphonocryptida n. sp., is described from Laos based on scanning electron microscopy, micro-computed tomography and molecular barcoding. The species of Dawydoffia are among the shortest and widest of the Polyzoniida, and resemble in habitus those of the Siphonocryptida genus Siphonocryptus Pocock, 1894, both having pleurites that are completely fused to the tergites. Dawydoffia was previously placed in the family Siphonotidae Cook, 1895, but can be identified as a member of the Holarctic Hirudisomatidae Silvestri, 1896 based on the following morphological characters: the uplifted posterior margins of the tergites, the collum covering part of the head, the position of the male gonapophysis or pseudopenis, the retracted telson, and the ozopores situated close to the tergal margin. However, both Dawydoffia species have a slender paronychium, a character previously known only from the Siphonotidae, but also documented here for Hirudisoma roseum (Victor, 1839). A slight redefinition of the Polyzoniida families is provided. The two Dawydoffia species differ in their coloration, as well as in their somatic and gonopodal characters.

Integrative redescription of the enigmatic monotypic alpine pill millipede genus Simplomeris Verhoeff, 1936 (Glomerida, Glomeridae, Haploglomerinae).

The European pill millipedes (Glomerida) are especially rich in species and genera of enigmatic status, which have been neither reviewed nor revised since more than 70 years. One of these genera is Simplomeris Verhoeff, 1936 with its single species S. montivaga (Faës, 1902), a high-altitude endemic only known from <5 collection events in the Simplon Valley and the Aletsch Glacier, southern Switzerland. Being one of only a handful of genera in the subfamily Haploglomerinae, distributed in Europe and SE Asia, its morphology is of greater interest in a future phylogenetic sorting of the Glomerida. Fresh material, which was encountered by the author 10 years ago from both known localities, allows here a redescription of this rarely encountered genus. Aside from the first photograph of a living specimen, scanning electron microscopy and genetic barcoding of the COI gene have been conducted. Morphologically, the telopods of Simplomeris resemble those of Haploglomeris Verhoeff, 1906 from the eastern Alps and confirm the present position in the Haploglomerinae. Genetically, Simplomeris belongs to the Glomeris klugii Brandt, 1833 species-group and is particularly close to high-altitude endemics from the southwestern and Bergamasque Alps, especially G. primordialis Verhoeff, 1930, G. transalpina Koch, 1836, and G. oropensis Verhoeff, 1934. Genetic barcoding data confirm the three colour varieties of S. montivaga, S. montivaga var. montivaga Verhoeff, 1936 syn. nov., S. montivaga var. simplonensis Verhoeff, 1936: syn. nov., S. montivaga var. berisalensis Verhoeff, 1936 syn. nov. to be just that, and all three are formally synonymized under S. montivaga.

New Thai giant pill-millipede species, with new genetic barcoding data (Diplopoda, Sphaerotheriida, Zephroniidae).

Three new species of giant pill-millipedes, Sphaerobelum meridionalis Bhansali Wesener sp. nov., Zephronia chrysomallos Bhansali Wesener sp. nov. and Zephronia erawani Bhansali Wesener sp. nov. are described based on museum samples from Thailand. All three species are described in an integrative manner, combining light microscopy, scanning electron microscopy, CT scans and genetic barcoding. Genetic barcoding was successfully conducted for all holotypes of the new species. In addition, genetic barcoding data of four recently described Thai Zephronia species, Zephronia lannaensis Likhitrakarn Golovatch, 2021 in Likhitrakarn et al. 2021, Z. phrain Likhitrakarn Golovatch, 2021, Z. panhai Srisonchai et al. 2021 and Z. golovatchi Srisonchai et al. 2021, together with new locality records, were added to the dataset. Our dataset includes all published sequences of the family Zephroniidae, including all but one (Z. enghoffi Srisonchai et al., 2021) of the described species from Thailand, Laos and Cambodia. All new species are genetically distant from other Zephroniidae from Thailand and surrounding countries showing uncorrected p-distances of >10 %. S. meridionalis sp. nov. is genetically and morphologically close to a recently described aberrant Sphaerobelum, S. aesculus Rosenmejer Wesener, 2021, as well as an unspecified specimen from Malaysia, and might represent a genus different from Sphaerobelum Verhoeff, 1924. Both new Zephronia species are geographically, morphologically and genetically close to Z. panhai, but differ from the latter by >10% p-distance in the COI gene and numerous morphological characters. Virtual cybertypes of the holotype of Zephronia erawani sp. nov. and of a paratype of Z. chrysomallos sp. nov. were created and made publicly accessible.

First record of the order Polyzoniida from the Indian subcontinent with an integrative description of a new genus (Diplopoda, Colobognatha, Siphonotidae).

Considering the large area of the Indian subcontinent, its known millipede diversity is sparse with only ca. 270 described species in 90 genera, 25 families and 11 orders. So far, not a single polyzoniidan millipede has been described from India. The order Polyzoniida Cook, 1895 is one of the most species-poor millipede groups with less than 80 described species, and includes taxonomically problematic groups, especially in the family Siphonotidae Cook, 1895. Here we report the first representatives of the order Polyzoniida from India, and describe the new genus Theratta n. gen., with the three species Theratta mannavan n. sp., Theratta eravikulam n. sp., and Theratta shola n. sp., using scanning electron microscopy and COI genetic barcoding. Based on the morphological characters we place Theratta n. gen., and the three new species in the tribe Rhinotini Hoffman, 1977 of the family Siphonotidae. The gonopods of the new genus are similar to those of the genus Rhinotus Cook, 1896, but differ from all other Rhinotini in the modification of the podomere II of the anterior telopod, carrying a mesal process on the posterior side and a lamellar process on the anterior side. The three newly described species show a high interspecific genetic distance of 20.925.3%. The specimens were collected from the Shola forests (high altitude montane forests) of the southern Western Ghats in Kerala, India. We suggest that more extensive sampling in this area and in the Indian subcontinent in general will yield more new millipede species for science and expand our understanding of the hitherto neglected Indian millipede fauna.

A previously unknown feeding mode in millipedes and the convergence of fluid feeding across arthropods.

We report fluid feeding with a sucking pump in the arthropod class Diplopoda, using a combination of synchrotron tomography, histology, electron microscopy, and three-dimensional reconstructions. Within the head of nine species of the enigmatic Colobognatha, we found a pumping chamber, which acts as positive displacement pump and is notably similar to that of insects, showing even fine structural convergences. The sucking pump of these millipedes works together with protractible mouthparts and externally secreted saliva for the acquisition of liquid food. Fluid feeding is one of the great evolutionary innovations of terrestrial arthropods, and our study suggests that it evolved with similar biomechanical solutions convergent across all major arthropod taxa. While fluid-feeding insects are megadiverse today, it remains unclear why other lineages, such as Colobognatha, are comparably species poor.

The millipede family Striariidae Bollman, 1893. VII. Petra sierwaldae, n. gen., n. sp., a minute millipede from Idaho, USA (Diplopoda, Chordeumatida, Striariidae).

Petra sierwaldae, n. gen., n. sp. is described from males and females collected at four localities in Kootenai County, Idaho, USA. The new genus is defined by its unique gonopod and ninth leg anatomy, a notched collum and single ommatidium on each side of the head, and lacking third coxal flasks as well as having modified fifth coxae.

How to uncoil your partner--"mating songs" in giant pill-millipedes (Diplopoda: Sphaerotheriida).

The stridulation of the giant pill-millipede genus Sphaerotherium from South Africa, one of only three groups of millipedes that produce sounds, was studied. One hundred one stridulation series of a total of nine different species (Sphaerotherium dorsaloide, Sphaerotherium hanstroemi, Sphaerotherium mahaium, Sphaerotherium similare, Sphaerotherium punctulatum, Sphaerotherium convexitarsum, Sphaerotherium dorsale, Sphaerotherium rotundatum, and Sphaerotherium perbrincki) were analyzed. Stridulation sounds are produced only with a special field of ribs on the posterior surface of the posterior telopod, which is actively moved over a field of sclerotized nubs on the inner margin of the anal shield. The Sphaerotherium male usually stridulates only when in contact with a female to initiate mating. This seems to prevent the female from volvating into a ball or stimulate the female to uncoil when already rolled in. The sound analyzes revealed a broad frequency spectrum in all stridulation sounds produced, without obvious differences in frequency distribution among species. However, the temporal pattern of the stridulation varies greatly between species and seems to be species-specific, arguing for a species recognition function of the stridulation during courtship behavior. A single species (S. punctulatum) was found to stridulate during mating while three species also show postcopulatory stridulation. Apparently, pill-millipedes are not capable of acoustic perception, as no hearing organs are known, indicating that the communication is mainly based on perception of temporal vibration patterns, and not of the acoustic signal itself. The need to overcome the rolling-in reflex of the female is developed as a hypothesis why stridulation exists only in millipedes able to coil into a ball, and apparently evolved four times independently in the superorder Oniscomorpha.

Description of the first species of Glomeridesmida from Thailand (Diplopoda, Glomeridesmida, Glomeridesmidae).

With three genera and 35 previously known species from India, SE Asia, Central and South America, Glomeridesmida are one of the least diverse Diplopoda groups. Here we describe Glomeridesmus siamensis sp. nov., the first species of the order Glomeridesmida from Thailand. The geographically nearest confamiliar species have been described from southern India, Sumatra and Java. The species is described combining photographs, light- and scanning electron microscopy of mature and younger males, females and juveniles. Several characters are illustrated for the first time for an Asian representative of the family Glomeridesmidae. In addition to the type locality of G. siamensis sp. nov. from Krabi province, locality data of unidentified Glomeridesmus from Thailand are also given. These data are providing further evidence that the Glomeridesmida are not uncommon, but overlooked as they are small and difficult to collect. The unusual telopods and other morphological characters of G. siamensis sp. nov. differ considerably from the few Glomeridesmus males described from Central and South America as well as from India, but the unclear status of two generic names available for species from Indonesia prevents us from adding another generic name to this small and understudied order.

Is it time to describe new species without diagnoses?A comment on Sharkey et al. (2021).

New methods in taxonomy and systematics can influence the overall practice of formally naming and describing biodiversity. DNA barcoding has been controversial since its emergence, but now, large scale species descriptions exclusively based on barcodes have created what can be called a 'new quality of performance. Its limitations are discussed from different perspectives: nomenclature, general pragmatism, and problems of DNA-based species delimitation in the light of the central aim of achieving a robust and stable nomenclature of organisms, essential for all applications of biodiversity research. This issue needs to be addressed to prevent restraining the progress of taxonomy and its ability to contribute to modern science.

The origins of the giant pill-millipedes from Madagascar (Diplopoda: Sphaerotheriida: Arthrosphaeridae).

Giant pill-millipedes (order Sphaerotheriida) are large-bodied millipedes without poison glands which can roll-up into a complete ball. Their disconnected area of distribution spanning South Africa, Madagascar, India, SE Asia, Australia and New Zealand makes them interesting model organisms for biogeographic studies. The here presented phylogeny is based on a molecular dataset covering all areas of distribution with a special focus on Madagascar, where some species of giant pill-millipedes show island gigantism, reaching the size of a baseball. For our study, two mitochondrial genes (partial 16S rRNA and COI) as well as the complete nuclear 18S rDNA were sequenced. While many recent vertebrate studies hint that the ancestors of the recent Malagasy fauna crossed the >350 km wide Mozambique Channel several times, no such crossing was discovered in the Sphaerotheriida. For the first time in a molecular phylogenetic study of soil arthropods, a Madagascar-India group, the family Arthrosphaeridae, is recovered, hinting to a Gondwanan origin of the Sphaerotheriida. The Malagasy-Indian family Arthrosphaeridae forms a monophyletic, statistically well-supported group in all obtained trees. The giant pill-millipedes from Madagascar are paraphyletic because the Malagasy genus Sphaeromimus is the sister-taxon of the Indian Arthrosphaera. In Sphaeromimus, an ecotone shift occurred only once: the spiny forest species Sphaeromimus musicus forms the sister-clade to the species collected in rainforests and littoral rainforests. The two species of the Malagasy genus Zoosphaerium which express island gigantism form a monophyletic group in some trees, but these trees lack good statistical support. Deeper nodes inside the Sphaerotheriida, like the position of the Australian genera Procyliosoma and Epicyliosoma, the Southeast Asian family Zephroniidae and the South African genus Sphaerotherium could not be resolved. This study is the first genetic study inside the order Sphaerotheriida and provides a proper basis for future molecular biogeographic studies in millipedes and soil organisms from Madagascar.

Ecotone shifts in southern Madagascar: first barcoding data and six new species of the endemic millipede genus Riotintobolus (Spirobolida, Pachybolidae).

Six new species of the Spirobolida millipede genus Riotintobolus Wesener, 2009, are described from the spiny forest in southern Madagascar utilising genetic barcoding, drawings and scanning electron microscopy: Riotintobolus tsimelahy sp. nov., R. mangatsiaka sp. nov., R. lavanono sp. nov., R. bovinus sp. nov., R. antafoky sp. nov. and R. makayi sp. nov. One other Riotintobolus population from the spiny forest might represent an additional species based on genetic data, but it cannot be described as no male specimens were collected. At present, the genus Riotintobolus Wesener, 2009 has eight species from the spiny forest and two species from the littoral rainforest. A determination key to all ten species of the genus is provided. Molecular data reveal that the two critically endangered species from the humid littoral rainforest are not closely related to one another, but have their closest relative in the dry spiny forest ecosystem. Riotintobolus mandenensis Wesener, 2009, only known from the southern littoral rainforest of Mandena is related to R. tsimelahy sp. nov. from the nearby spiny forest at Tsimelahy with a p-distance of 11%, while R. minutus Wesener, 2009 from the littoral forest of Sainte Luce is more distant to all other Riotintobolus species, but more closely related to R. bovinus sp. nov. from the southwestern forest of the Makay.

The millipedes collected by the Museum "La Specola" on Madagascar 1989/1991, with the description of three new species of giant pill-millipedes (Diplopoda, Sphaerotheriida, Arthrosphaeridae).

A large collection of millipedes (Diplopoda) from Madagascar, belonging to the Museum "La Specola" in Florence, Italy were investigated. The collection includes three new species of the giant pill-millipede genus Zoosphaerium Pocock, 1895 which are described here as Zoosphaerium mangabe Wesener, sp. nov., Z. bartolozzii Anilkumar & Wesener, sp. nov., and Z. taitii Anilkumar & Wesener, sp. nov., all belonging to the Z. coquerelianum species group. The latter two are currently only known from a single site. Other specimens belonging to eight orders (Polyxenida, Sphaerotheriida, Polyzoniida, Siphonophorida, Chordeumatida, Polydesmida, Spirobolida, and Spirostreptida) are listed. Three tropical tramp species, Pseudospirobolellus avernus (Butler, 1876), Glyphiulus granulatus Gervais, 1847, and Chondromorpha xanthotricha (Attems, 1898) are recorded for the first time from Madagascar. New locality data is provided for Zoosphaerium neptunus (Butler, 1872), Z. villosum Wesener & Sierwald, 2005, Z. blandum (de Saussure & Zehntner, 1897), Sphaeromimus musicus (de Saussure & Zehntner, 1897), Rhinotus purpureus (Pocock, 1894), Hylekobolus andasibensis Wesener, 2009, Aphistogoniulus infernalis Wesener, 2009, Ostinobolus rufus Wesener, 2009, Ostinobolus subterraneus Wesener, 2009, Dactylobolus bivirgatus (Karsch, 1881), and Eumekius antimena (de Saussure & Zehntner, 1901).

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.

The millipede family Polyxenidae (Diplopoda, Polyxenida) in the faunas of the Crimean Peninsula and Caucasus, with notes on other European Polyxenidae.

The Polyxenidae in the fauna of the Crimeo-Caucasian region is represented by four species: Polyxenus lagurus (Linnaeus, 1758) (= P. lagurus caucasicus Lignau, 1924, syn. n.), Propolyxenus argentifer (Verhoeff, 1921) comb. n. (= P. trivittatus Verhoeff, 1941, = P. sokolowi Lignau, 1924, both syn. n.), a new species, Polyxenus lankaranensis sp. n., and an undescribed Polyxenus sp. The distributions of all these species in the region concerned are mapped, based on old and new records. A molecular phylogeny based on COI sequences is used to study the relationship within and among the genera Polyxenus and Propolyxenus from Western Europe to the southern Caucasus. The results highlight the presence of a number of undescribed species of Polyxenus across this region.

A first phylogenetic analysis of Giant Pill-Millipedes (Diplopoda: Sphaerotheriida), a new model Gondwanan taxon, with special emphasis on island gigantism.

The phylogeny of the Giant Pill-Millipedes, order Sphaerotheriida, is investigated using a new morphological character matrix comprising 89 characters. The majority of these characters are employed for the first time in millipedes. All trees obtained agree on the monophyletic status of the Sphaerotheriida and several of its tribes, each restricted to a modern land mass. The species from Madagascar displaying island gigantism do not form a monophyletic group. The classic division of Giant Pill-Millipedes into two families, Sphaerotheriidae and Zephronidae, was not reflected in the analysis. The genus Procyliosoma is the sister-group to all other Sphaerotheriida, rendering the family Sphaerotheriidae paraphyletic. A new family-level classification of Giant Pill-Millipedes, based on the current phylogeny, is introduced. The new family Procyliosomatidae contains only the genus Procyliosoma, distributed in Australia and New Zealand. The family Zephronidae remains unchanged, while the family Sphaerotheriidae now incorporates only the African Giant Pill-Millipede genera. All genera from southern India and Madagascar form a monophyletic group and are placed in the new family Arthrosphaeridae. The Malagasy genus Sphaeromimus is more closely related to the Indian Arthrosphaera species than to other genera from Madagascar. A biogeographical analysis identifies the group as a Gondwana taxon (with a notable absence from South America). The current phylogeny of Giant Pill-Millipede families mirrors perfectly the suggested break-up of Gondwana fragments 160-90 Ma. No evidence for a dispersal event could be found, highlighting the importance of Giant Pill-Millipedes as a potential model taxon.

First records of giant pill-millipedes from Laos (Diplopoda, Sphaerotheriida, Zephroniidae).

Before this study, giant pill-millipedes (order Sphaerotheriida) were unknown from Laos despite their presence in all surrounding countries. As by-catch from collections by arachnologists, 31 specimens of Sphaerotheriida from Laos became available for study. The sample included 14 species. Three species were only represented by females, which are described but cannot be named. Of the remaining 11 species, a single species belongs to the genus Zephronia Gray, 1832: Z. laotica n. sp.; and the other ten belong to the genus Sphaerobelum Verhoeff, 1924: S. bolavensis n. sp., S. phouloei n. sp., S. denticulatum n. sp., S. spinatum n. sp., S. lachneeis n. sp., S. peterjaegeri n. sp., S. nigrum n. sp., S. splendidum n. sp., S. laoticum n. sp., and S. schwendingeri n. sp. This more than doubles the known diversity of Sphaerobelum. Here, I integratively describe these species, combining morphology and DNA barcodes with a molecular analysis including all Zephroniidae species deposited on GenBank-including the only giant pill-millipede species known from Cambodia, Zephronia dawydoffi Attems, 1953. An updated determination key to the species of the genus is presented. Zephronia laotica n. sp. belongs to the monophyletic Zephronia sensu stricto group, which is confirmed by molecular barcoding. In contrast, most species of Sphaerobelum are in a weakly supported clade. Genetically, Sphaerobelum species differ greatly from one another, with most p-distances >15%. The lowest observed p-distance (9.8%) is between S. truncatum Wongthamwanich et al. 2012 from Thailand and S. peterjaegeri n. sp.