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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

Dwarfs under dinosaur legs: a new millipede of the order Callipodida (Diplopoda) from Cretaceous amber of Burma.

The entire Mesozoic Era is rather poor in millipede (class Diplopoda) fossils, with less than a dozen species being taxonomically described. Here, we describe the first fossil millipede of the order Callipodida, Burmanopetaluminexpectatum gen. nov. et sp. nov., found in early Cenomanian amber of Burma, 98.79±0.62 Mya. The species possesses a number of morphological traits that exclude it from all extant suborders, and Burmanopetalidea suborder nov. and Burmanopetalidae fam. nov. are here erected to accommodate it. The new suborder can be recognized by the following unique characters: pleurotergal setae absent; telson with a specific spatulate shape twice the size of the penultimate body ring; hypoproct devoid of setae; and eyes composed of five well-separated ommatidia. While the callipodidan habitus seems to have remained generally unchanged for at least 99 million years, pleurotergal and hypoproctal setation, as well as the complexity of eyes in ground-dwelling forms may have evolved recently in the order. As B.inexpectatum gen. nov. et sp. nov. is the first true callipodidan in the fossil record, the minimum age of Callipodida is thus at least 99 Mya.

The Myriapoda of Halimun-Salak National Park (Java, Indonesia): overview and faunal composition.

The myriapod fauna of the mega-diverse country of Indonesia is so far insufficiently known, with no species lists or determination keys. In order to obtain an overview of the faunal composition of the Myriapoda in an Indonesian forest system, the fauna of the Halimun-Salak National Park in western Java was explored during the dry season (September-October 2015) in the framework of the German-Indonesian INDOBIOSYS project (Indonesian Biodiversity Discovery and Information System). A total of 980 Myriapoda specimens were collected by hand by 3-4 researchers from three different sites in the national park, from which 796 specimens were determined to a higher taxonomic level (class, order, family) and 617 specimens were determined to morphospecies. Among these, 27 were Symphyla (4%) (excluded from further analyses), 226 Chilopoda (28%) and 543 Diplopoda (68%). The Scolopendromorpha (64% of all identified centipedes) and Polydesmida (69% of all identified Diplopoda) were the most represented orders in our samples. Twenty-four morphospecies of Chilopoda were determined: one each of Scutigeromorpha and Lithobiomorpha, six Scolopendromorpha and sixteen Geophilomorpha. Nine orders of diplopods were present, with a total of 47 morphospecies: one each of Polyxenida, Glomeridesmida and Chordeumatida, two each of Glomerida, Spirobolida and Siphonophorida, seven of Sphaerotheriida, ten of Spirostreptida and 21 of Polydesmida. Two species curves were obtained to have a first idea about the myriapod diversity in the Halimun-Salak National Park and to compare the three individual collecting sites. Our results depict the Scolopendromorpha as the most common centipedes in Javanese rainforests and the Geophilomorpha as the most species-rich order. In contrast, the Polydesmida were the most dominant millipede group with 167 specimens and with 13 morphospecies the family Paradoxosomatidae was the most diverse.

Heat tolerance and acclimation capacity in subterranean arthropods living under common and stable thermal conditions.

Cave-dwelling ectotherms, which have evolved for millions of years under stable thermal conditions, could be expected to have adjusted their physiological limits to the narrow range of temperatures they experience and to be highly vulnerable to global warming. However, most of the few existing studies on thermal tolerance in subterranean invertebrates highlight that despite the fact that they show lower heat tolerance than most surface-dwelling species, their upper thermal limits are generally not adjusted to ambient temperature. The question remains to what extent this pattern is common across subterranean invertebrates. We studied basal heat tolerance and its plasticity in four species of distant arthropod groups (Coleoptera, Diplopoda, and Collembola) with different evolutionary histories but under similar selection pressures, as they have been exposed to the same constant environmental conditions for a long time. Adults were exposed at different temperatures for 1 week to determine upper lethal temperatures. Then, individuals from previous sublethal treatments were transferred to a higher temperature to determine acclimation capacity. Upper lethal temperatures of three of the studied species were similar to those reported for other subterranean species (between 20 and 25°C) and widely exceeded the cave temperature (13-14°C). The diplopod species showed the highest long-term heat tolerance detected so far for a troglobiont (i.e., obligate subterranean) species (median lethal temperature after 7 days exposure: 28°C) and a positive acclimation response. Our results agree with previous studies showing that heat tolerance in subterranean species is not determined by environmental conditions. Thus, subterranean species, even those living under similar climatic conditions, might be differently affected by global warming.

Four new species of giant pill-millipedes from Vietnam (Sphaerotheriida, Zephroniidae).

Four new species of giant pill-millipedes (Sphaerotheriida) are described from southern Vietnam: Sphaerobelum cattiense sp. nov., Sphaerobelum konkakinhense sp. nov., Zephronia konkakinhensis sp. nov. and Zephronia montis sp. nov. An updated generic diagnosis of Sphaerobelum Verhoeff, 1924 and a key to its species are provided.

An integrative and citizen science based approach to the rediscovery and redescription of the only known high-altitude endemic Pill Millipede, Glomeris aurita Koch (Diplopoda, Glomerida).

The pill millipede species Glomeris aurita Koch, 1847 remained of relative unknown origin and appearance until its recent rediscovery in samples from the Bergamasque Alps, northern Italy. In order to provide an integrative redescription and accurate identification of the high-altitude microendemic G. aurita, COI barcode sequences from three individuals coming from two different localities were obtained. These sequences are compared with those of the syntopic endemic G. oblongoguttata Verhoeff, 1894, the widespread black morph of G. romana Verhoeff, 1900, as well as several widespread species including G. marginata Villers, 1789, G. connexa Koch, 1847, and G. klugii Brandt, 1833, which have rare colour morphs that exhibit some similarity to G. aurita. To rule-out any identity confusion of G. aurita with other high-altitude or little-known Italian Glomeris, specimens of G. transalpina Koch, 1836, G. oropensis Verhoeff, 1934, and G. primordialis Verhoeff, 1932 were also added to the dataset. Altogether, 24 sequences were compared. Morphologically, the specimens of G. aurita were studied utilizing scanning electron microscopy as well as non-invasive micro-CT technology. The distribution of both Bergamasque endemics, G. aurita and G. oblongoguttata, could be mapped and compared utilizing samples from the Museo civico di Scienze Naturali di Bergamo, as well as photographic evidence from an Italian naturalist forum. G. aurita has a very short active period and is the first known pill millipede species restricted to mountain tops and cold places, possibly representing a Nunatak survivor.