Search

The family Dalodesmidae Cook, 1896 in the fauna of Madagascar is reviewed and shown to presently encompass eight species in three genera: Dalodesmus Cook, 1896 (six species), Eutubercularium Brölemann, 1916 (one species), and Phymatodesmus de Saussure & Zehntner, 1897 (one species). These genera are diagnosed, and their respective species keyed, all being endemic to Madagascar proper and/or the immediately adjacent islets of Nosy Be and/or Nosy Sakatia. Dalodesmus currently contains six species, including two new, all supplied either with brief descriptive notes and available iconography or extensive descriptions and new illustrations, as follows: D.hamatus (Brandt, 1841), from an unspecified locality in Madagascar, now redescribed from a male specimen from Makira, northeastern Madagascar; D.odontopezus (Attems, 1898), from Nosy Be Isle; D.orator Hoffman, 1974, from Ambohimitombo, central Madagascar; D.tectus Cook, 1896 (= D.hova (de Saussure & Zehntner, 1897), syn. nov. ), originally described from an unspecified locality in central Madagascar, with a male type and additional males identified as D.hova recorded from the Andasibe National Park (= Périnet) in east-central Madagascar. Additionally, two new species are described, D.speophilus sp. nov. , from Grotte de Anjohibe, northwestern Madagascar, and D.kompantsevi sp. nov. , from Montagne d'Ambre, northern Madagascar. Both the type species of Dalodesmus and its synonym Tubercularium Attems, 1898, D.tectus and D.odontopezus , respectively, are partly revised and illustrated based on holotypes, while D.hamatus is duly described and illustrated based on the first discovery of a male. Both Eutubercularium (Mesibov, Wesener & Hollier, 2018), from Nosy Be Isle, and Phymatodesmus are still monospecific, comprising only E.voeltzkowi (Mesibov, Wesener & Hollier, 2018), from Nosy Be Isle, and P.sakalava (de Saussure & Zehntner, 1901), from Andasibe, respectively. The latter species is fully redescribed and illustrated for the first time from male material., Competing Interests: The authors have declared that no competing interests exist., (Thomas Wesener, Nesrine Akkari, Sergei I. Golovatch.)

Among the more than 80 species of the common pill millipede genus Glomeris Latreille, 1803, there are several microendemic species that have not been recorded for the last 80-120 years. To discover whether these species are colour morphs of widespread species or true local endemics is important from a conservation point of view as well as for understanding the biogeography and evolution of the group. The author received three specimens that were morphologically identical to C. L. Koch's 175-year-old first description of Glomeris cingulata Koch, 1847 from the Triglav Mountain in Slovenia, close to the border with Italy. No clear specimen-based records are available for G. cingulata and the type specimen is apparently lost. In order to clarify the taxonomy of this microendemic species, an integrative redescription was conducted, including scanning electron microscopy and DNA barcoding. Glomeris cingulata is a distinct species, with genetic distances of 12.6-15.5% compared to the seven syntopic and numerous other widespread Glomeris species. Based on characters of the first description and following other authors, the synonymy of Glomeris cingulata intercedens Latzel, 1884 under Glomeris transalpina Koch, 1836 is confirmed. The dark colour with posterior red bands closely resembles that of some other high-altitude Glomeris species like G. transalpina Koch, 1836, Glomeris aurita Koch, 1847 and Glomeris oropensis Verhoeff, 1936. Glomeris cingulata is genetically close to, but distant enough from the small-bodied and widespread taxa like Glomeris pustulata Latreille, 1804 and Glomeris tetrasticha Brandt, 1833.

Based mainly on the results of generalized American biodiversity inventory programmes, several samples of Malagasy Spirobolida millipedes became available to study. Based on this material, five new species of Spirobolida are described from Madagascar: four potential microendemic species, Aphistogoniulus amberivery sp. nov. from the Amberivery forest, A. manombo sp. nov., from the lowland rainforest of Manombo, Spiromimus endemicus sp. nov. from the Montagne des Français, and a species from the littoral forest of Tampolo, tentatively placed in the genus Eucarlia Brölemann, 1913, E. tampolo sp. nov.. A fifth species appears to be more widespread and is a morphologically unusual species of the genus Alluviobolus Wesener, 2009, A. omega sp. nov.. Aphistogoniulus manombo sp. nov. was previously described as a population of A. jeekeli Decker & Wesener, 2011. One of the most enigmatic Malagasy millipede species, Spirobolus olympiacus Karsch, 1881, is redescribed based on topotypic material as Colossobolus olympiacus (Karsch, 1881) new combination. Additional locality data is provided for 14 other Spirobolida species, of which 11 are listed on the IUCN Red List: Aphistogoniulus aridus Wesener, 2009, A. diabolicus Wesener, 2009, A. erythrocephalus (Pocock, 1893), A. hova (de Saussure & Zehntner, 1897), Colossobolus semicyclus Wesener, 2009, Spiromimus albipes Wesener & Enghoff, 2009, S. litoralis Wesener & Enghoff, 2009, S. simplex Wesener & Enghoff, 2009, S. triaureus Wesener & Enghoff, 2009, S. univirgatus deSaussure & Zehntner, 1901, Flagellobolus pauliani Wesener, 2009, Dactylobolus bivirgatus (Karsch, 1881), Madabolus maximus Wesener & Enghoff, 2008 and Hylekobolus rufus Wesener, 2009. Fourteen new genetic barcoding COI sequences are provided for nine species: Hylekobolus rufus, Colossobolus semicyclus, Aphistogoniulus amberivery sp. nov., A. aridus, A. diabolicus, A. erythrocephalus, A. hova, Spiromimus univirgatus and S. scapularis Wesener & Enghoff, 2009.

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., Competing Interests: The authors have declared that no competing interests exist., (Ruttapon Srisonchai, Natdanai Likhitrakarn, Chirasak Sutcharit, Thomas Wesener.)

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.

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.

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.

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.

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.

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.

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.

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.

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.

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., (Thomas Wesener, Nattarin Wongthamwanich, Leif Moritz.)

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. 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. R. antafoky Wesener, 2009, only known from the southern littoral rainforest of Mandena is related to sp. nov. and R. makayi sp. nov. Wesener, 2009 from the littoral forest of Sainte Luce is more distant to all 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., (Thomas Wesener.)

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 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)., (Thomas Wesener, Pooja Avinipully Anilkumar.)

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., (Irina Semenyuk, Sergei I. Golovatch, Thomas Wesener.)

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., Competing Interests: The authors have no competing interest to declare., (© 2019 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

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 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.

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.

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., Competing Interests: The authors declare there are no competing interests.

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.

Polydesmus (Pterodesmus) voeltzkowi nom. nov. is proposed as a replacement name for Polydesmus (Pterodesmus) sakalava de Saussure Zehntner, 1901, now Dalodesmus sakalava (de Saussure Zehntner, 1901), a junior primary homonym of Polydesmus sakalava de Saussure Zehntner, 1897.

The Black Pill Millipede, Glomerismarginata , is the best studied millipede species and a model organism for Diplopoda. Glomerismarginata is widespread, with numerous colour morphs occurring across its range, especially in the south. This study investigates whether colour morphs might represent cryptic species as well as the haplotype diversity and biogeography of G.marginata . The results of the COI barcoding fragment analysis include 97 G.marginata , as well as 21 specimens from seven potentially related species: G.intermedia Latzel, 1884, G.klugii Brandt, 1833 ( G.undulata C.L. Koch, 1844), G.connexa Koch, 1847, G.hexasticha Brandt, 1833, G.maerens Attems, 1927, G.annulata Brandt, 1833 and G.apuana Verhoeff, 1911. The majority of the barcoding data was obtained through the German Barcode of Life project (GBOL). Interspecifically, G.marginata is separated from its congeners by a minimum uncorrected genetic distance of 12.9 %, confirming its monophyly. Uncorrected intraspecific distances of G.marginata are comparable to those of other widespread Glomeris species, varying between 0-4.7%, with the largest genetic distances (>2.5 %) found at the Mediterranean coast. 97 sampled specimens of G.marginata yielded 47 different haplotypes, with identical haplotypes occurring at large distances from one another, and different haplotypes being present in populations occurring in close proximity. The highest number of haplotypes was found in the best-sampled area, western Germany. The English haplotype is identical to northern Spain; specimens from southern Spain are closer to French Mediterranean specimens. Analyses (CHAO1) show that approximately 400 different haplotypes can be expected in G.marginata . To cover all haplotypes, it is projected that up to 6,000 specimens would need to be sequenced, highlighting the impossibility of covering the whole genetic diversity in barcoding attempts of immobile soil arthropod species.

The presence of a swinging tentorium is a key apomorphy of Myriapoda, but this character has been studied in detail in only few species. Here the tentorium, i.e., the peristomatic skeleton of the preoral chamber, is comparatively studied in three species of the millipede order Sphaerotheriida Brandt, 1833. Since dissections of the fragile tentorial components proved to be difficult, despite the large head size, they were analysed mainly in situ via micro-computed tomography. Our results confirm previous observations of large differences in the tentorial construction in the giant pill-millipedes compared to chilognathan diplopods. The tentorium of Sphaerotheriida consists of a curved, plate-like epipharyngeal bar with distal projections, an elongate and thin hypopharyngeal bar, and a plate-like triangular posterior process; a transverse bar is absent. Only seven muscles attach at the tentorium in giant pill-millipedes, including two antennal muscles and two muscles of the gnathochilarium. Within the order Sphaerotheriida, the composition of the tentorium and its muscular equipment seems to be conserved, except for some variability in the shape of the epipharyngeal bar. As the transverse bar has been considered essential for the mobility of the tentorium in myriapods, its absence in Sphaerotheriida may indicate that their tentorium is not capable of performing a swing. Loss of tentorial mobility may also pertain to the order Glomerida Brandt, 1833, inferred here from the absence of a posterior process. An apparently immobile tentorium in Glomerida and Sphaerotheriida can straightforwardly be correlated with transformations of the head related to their ability of volvation. The different transformations of the tentorium, here hypothesised to cause immobility, may support current assumptions that the ability of volvation evolved convergently in Glomerida and Sphaerotheriida. This conclusion, however, still requires more detailed studies of the head anatomy in Glomerida and Glomeridesmida Cook, 1895.

The pill millipedes of the order Glomerida are a moderately diverse group with a classical Holarctic distribution pattern. Their classification is based on a typological system utilizing mainly a single character complex, the male telopods. In order to infer the apomorphies of the Glomerida, to elucidate its position in the Pentazonia, and to test the monophyly of its families and subfamilies, we conduct the first phylogenetic analysis of the order. To provide additional characters, we comparatively analyze the mandible using scanning electron microscopy. The final character matrix consists of 69 characters (11 mandible characters) and incorporates 22 species from 20 of the 34 pill millipede genera, representing all families and subfamilies, except the monotypic Mauriesiinae. Two species from each of the two other Pentazonian orders Sphaerotheriida and Glomeridesmida, as well as two Spirobolida, are included as outgroup taxa. The Glomerida are recovered as monophyletic and are supported by five apomorphies. Within the Pentazonia, the Glomeridesmida are recovered as the sister group to the classical Oniscomorpha (Sphaerotheriida + Glomerida) with weak support. The analysis provides little resolution within the Glomerida, resulting in numerous polytomies. Further morphological characters and/or the addition of molecular analyses are needed to produce a robust phylogenetic classification of the Glomerida., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Madagascar is one of the world's most important hotspots of biodiversity and a center for localized endemism. Among the highly endemic faunal elements are the giant pill-millipedes, order Sphaerotheriida, which are severely understudied in Madagascar. Here we provide descriptions of two new species of endemic giant-pill millipedes of the genus Zoosphaerium Pocock, 1895: Zoosphaerium bemanevika n. sp. and Zoosphaerium minutus n. sp.. Zoosphaerium bemanevika n. sp. belongs to the Z. coquerelianum species-group, while Z. minutus n. sp. is not assignable to a species-group. An updated key to the 19 species of the Z. coquerelianum group is provided. Zoosphaerium minutus n. sp. has a body length of <20 mm and thus is the smallest known member of the genus, being of similar size as the larger species of Microsphaerotherium Wesener & VandenSpiegel, 2007. Both species are described utilizing drawings, scanning electron microscopy and genetic barcoding based on the mitochondrial COI gene. MicroCT imaging is applied to study internal morphology non-destructively and in situ, allowing for a reconstruction of the head skeleton. Our results show that females of Z. bemanevika n. sp. exhibit island gigantism. The two newly described species are not closely related to one another but each to different species from the Marojejy Mountain and the lowland rainforest of the east coast. Both species occur sympatrically as microendemics in small patches of humid evergreen forest near Bemanevika in northern Madagascar, an only recently protected area that represents a Malagasy center of endemism.

Four new species of the chordeumatidan millipede genus Nepalella Shear, 1979, family Megalotylidae, are described from caves in southern China: N. troglodytes sp. nov., N. lobata sp. nov., N. jinfoshan sp. nov., and N. wangi sp. nov. Three of them (except N. lobata sp. nov.) are presumed troglobites. Additional locality records of two cave congeners, N. caeca Shear, 1999 and N. grandoides Golovatch, Geoffroy & Mauriès, 2006, are also provided. DNA-barcoding based on the COI mitochondrial gene is documented in this genus and for species of the order Chordeumatida outside Europe for the first time. Interspecific p-distances between Nepalella species amount to 8.5-15.9%, while intraspecific p-distances are 0-6.8%. The species of Nepalella are found to show a surprisingly low genetic distance from the European genus Atractosoma Fanzago, 1876, family Craspedosomatidae Gray in Jones, 1843, potentially based on the very limited number of barcoding sequences of the order Chordeumatida being available.

Laos has large areas of primary forest with a largely unexplored fauna. This is evidenced by millipedes, class Diplopoda, with fewer than 60 species being recorded from the country. In the widespread Southeast Asian "Star Millipede" genus Eutrichodesmus Silvestri, 1910 (family Haplodesmidae), only two of 49 recorded species have been found in Laos. Four new species of Star Millipedes are here described from caves in Laos: Eutrichodesmus steineri Liu & Wesener, sp. n. , Eutrichodesmus deporatus Liu & Wesener, sp. n. , Eutrichodesmus paraster Liu & Wesener, sp. n. and Eutrichodesmus parvus Liu & Wesener, sp. n. . A fifth species, for which only a female is available, remains unnamed. The defensive glands (ozopores) are found to be strongly or entirely suppressed in two of the new species, Eutrichodesmus deporatus Liu & Wesener, sp. n. and Eutrichodesmus paraster Liu & Wesener, sp. n. , both troglobionts, which is new to the family. All of the Star Millipedes were collected during Northern Lao-European Cave Project faunal surveys conducted by the Senckenberg Museum, Frankfurt. A key to the six species of Eutrichodesmus currently known to occur in Laos is provided.

Animal life in caves has fascinated researchers and the public alike because of the unusual and sometimes bizarre morphological adaptations observed in numerous troglobitic species. Despite their worldwide diversity, the adaptations of cave millipedes (Diplopoda) to a troglobitic lifestyle have rarely been examined. In this study, morphological characters were analyzed in species belonging to four different orders (Glomerida, Polydesmida, Chordeumatida, and Spirostreptida) and six different families (Glomeridae, Paradoxosomatidae, Polydesmidae, Haplodesmidae, Megalotylidae, and Cambalopsidae) that represent the taxonomic diversity of class Diplopoda. We focused on the recently discovered millipede fauna of caves in southern China. Thirty different characters were used to compare cave troglobites and epigean species within the same genera. A character matrix was created to analyze convergent evolution of cave adaptations. Males and females were analyzed independently to examine sex differences in cave adaptations. While 10 characters only occurred in a few phylogenetic groups, 20 characters were scored for in all families. Of these, four characters were discovered to have evolved convergently in all troglobitic millipedes. The characters that represented potential morphological cave adaptations in troglobitic species were: (1) a longer body; (2) a lighter body color; (3) elongation of the femora; and (4) elongation of the tarsi of walking legs. Surprisingly, female, but not male, antennae were more elongated in troglobites than in epigean species. Our study clearly shows that morphological adaptations have evolved convergently in different, unrelated millipede orders and families, most likely as a direct adaptation to cave life., Competing Interests: The authors have declared that no competing interests exist.

Sphaeropoeus hercules Brandt, 1833, the type species of one of the oldest genus names in Sphaerotheriida, is redescribed for the first time based on its fragmented type specimen. While damaged, the specimen shows characters clearly different from other, diverse and similar genera of the family Zephroniidae, such as Zephronia Gray, 1832 and Castanotherium Pocock, 1895. S. hercules has a record number of &gt;180 apical cones on the antennae, more than any other millipede. The species, together with another species of the genus Castanotherium, C. insigne (Brandt, 1833) is added to an existing character matrix currently comprising 100 characters and 46 species. A second never-revised species of Sphaeropoeus, S. variegatus Pocock, 1895, is also redescribed based on type material. The latter seems to belong to the genus Sphaeropoeus, but its provenance remains currently unknown. S. variegatus could not be added to the character matrix because several important characters could not be studied. The phylogenetic analysis resolved the position of Sphaeropoeus in the Zephroniidae, but gave no resolution of the intrafamiliar relationships of the family's numerous genera.

Local endemic species with their unique evolutionary history always stirred the interest of scientists. One such area especially rich in endemics is northern Italy. In case of pill millipedes of the genus Glomeris Latreille, 1803, only a single species is found in northern Europe, while 22 country-endemics alone are known from Italy. Many of these endemics, however, have not been studied in several decades; therefore we aimed to determine whether this diversity is the result of overlooked synonymies or natural processes. A focus was placed on the local endemics that are in some aspects morphologically similar to the widespread and variable G. klugii Brandt, 1833. The local endemics Glomeris larii Verhoeff, 1921, G. primordialis Verhoeff, 1930, G. oblongoguttata Verhoeff, 1894, G. oropensis Verhoeff, 1936, G. transalpina Koch, 1836, G. romana Verhoeff, 1900, G. ligurica Latzel, 1884 and G. apuana Verhoeff, 1911 were included in a molecular analysis incorporating ribosomal nuclear (28S) and mitochondrial (COI) genes. Individuals were sequenced and compared to 31 specimens from 18 localities of G. klugii. The final dataset included 657 base pairs for 56 terminals in the COI, and 14 terminals with 1068 base pairs in the combined 28S and COI analysis. Our analysis shows intraspecific distances of up to 5% in the COI gene in G. klugii that are not strictly correlated to geography or color pattern. G. larii is discovered to be genetically and morphologically identical to G. klugii and is synonymised with the latter. Interspecific distances in our dataset vary between 6.7 to 15.9%, with the lowest (6.7-9.0%) between G. primordialis and G. klugii. Our analysis confirms the species status of the local endemics G. primordialis, G. oblongoguttata, G. oropensis, G. transalpina, G. ligurica and G. apuana. We also confirm the synonymy of G. undulata Koch, 1844 under G. klugii. G. genuensis Latzel, 1886 is indistinguishable from G. ligurica., Competing Interests: The authors have declared that no competing interests exist.

At present, the millipede fauna of India comprises over 270 nominate species or subspecies in at least 90 genera, 25 families and 11 orders. As complete a catalogue as possible is compiled of all species or subspecies of Diplopoda formally reported from India, some of which remain dubious. Additionally, a checklist is also compiled of millipedes that have erroneously been recorded in India. Given also several literature lacunae, as well as numerous taxonomic problems, the number of species and even genera cannot be claimed precise. The following new transfers are proposed: Delarthrum pumilum (Attems, 1944), comb. n. ex Dasypharkis Attems, 1936; Stemmiulus crassipes (Carl, 1941), S. insolitus (Carl, 1941), S. mulierosus (Carl, 1937), S. plumipes (Carl, 1941) and S. vagans (Carl, 1941), all comb. n. ex Diopsiulus Silvestri, 1897; Cryptocorypha riparia (Carl, 1932), comb. n. ex Archandrodesmus Carl, 1932. A new record of Stenobolus insularis Carl, 1918 is given from India.

In order to evaluate the diversity of Central European Myriapoda species in the course of the German Barcode of Life project, 61 cytochrome c oxidase I sequences of the genus Cryptops Leach, 1815, a centipede genus of the order Scolopendromorpha, were successfully sequenced and analyzed. One sequence of Scolopendra cingulata Latreille, 1829 and one of Theatops erythrocephalus Koch, 1847 were utilized as outgroups. Instead of the expected three species (Cryptops parisi Brolemann, 1920; Cryptops anomalans Newport, 1844; Cryptops hortensis (Donovan, 1810)), analyzed samples included eight to ten species. Of the eight clearly distinguishable morphospecies of Cryptops, five (Cryptops parisi; Cryptops croaticus Verhoeff, 1931; Cryptops anomalans; Cryptops umbricus Verhoeff, 1931; Cryptops hortensis) could be tentatively determined to species level, while a further three remain undetermined (one each from Germany, Austria and Croatia, and Slovenia). Cryptops croaticus is recorded for the first time from Austria. A single specimen (previously suspected as being Cryptops anomalans), was redetermined as Cryptops umbricus Verhoeff, 1931, a first record for Germany. All analyzed Cryptops species are monophyletic and show large genetic distances from one another (p-distances of 13.7-22.2%). Clear barcoding gaps are present in lineages represented by >10 specimens, highlighting the usefulness of the barcoding method for evaluating species diversity in centipedes. German specimens formally assigned to Cryptops parisi are divided into three clades differing by 8.4-11.3% from one another; their intra-lineage genetic distance is much lower at 0-1.1%. The three clades are geographically separate, indicating that they might represent distinct species. Aside from Cryptops parisi, intraspecific distances of Cryptops spp. in Central Europe are low (<3.3%).

The Italian pill millipede species Glomeris apuana Verhoeff, 1911, is redescribed from fresh material and its COI barcoding fragment is sequenced. The new specimens were compared to the original type series, of which a lectotype was selected. G. apuana was apparently still viewed as a subspecies of G. ligurica, as its name cannot be found in 'Fauna Europaea', or any faunal lists or catalogues in the last 85 years. We show that the species is both genetically and morphologically unique. G. apuana is easy to identify based on its entirely black coloration in combination with the absence of any main striae on the thoracic shield. Genetically, G. apuana shows large p-distances of &gt;10% to four different populations of G. ligurica Latzel, 1884. G. apuana also differs from other sequenced Glomeris species, G. marginata Latreille, 1803, G. connexa Koch, 1847, and G. klugii Brandt, 1833 by p-distances of &gt;10%. Specimens of G. klugii from a population occurring in sympatry with G. apuana were newly sequenced. All records of G. apuana, a large, easy to identify and conspicuous species, are from a narrow coastal zone of the Apuan Alps, an area in which the species might be microendemic.

In order to evaluate the status of the only species of pill millipede (Glomerida) endemic to Germany, Glomeris malmivaga Verhoeff, 1912, a DNA barcoding study based on the COI mitochondrial gene was conducted. Sequences of G. malmivaga were compared to those of G. ornata Koch, 1847 from Slovenia, of which the former was previously described as a variety of the latter before being elevated to subspecies- and, recently, species-rank. Included in the analysis were specimens of G. helvetica Verhoeff, 1894, also originally described as a variety of G. ornata, which was supposed to be closely related to G. malmivaga based on its morphology, as well as geographical proximity of occurrence. Additionally, G. valesiaca Rothenbühler, 1899, which occurs in sympatry and looks quite similar to G. helvetica was also sequenced for the first time and included in the study. Sequences of four widespread Glomeris species, all occurring in close proximity to G. malmivaga, G. marginata Villers, 1789, G. connexa Koch, 1847, G. klugii Brandt, 1833 and G. intermedia Latzel, 1884 were downloaded from Genbank and incorporated in the analysis. While G. helvetica and G. valesiaca were found to be clearly separate from G. ornata (11.8-14.6% p-distance), G. malmivaga is almost identical to the latter (0.5% p-distance), despite the large geographical distance between both species. Because of their great morphological and genetical similarity, G. malmivaga n. syn. is synonymised under G. ornata.

Eupeyerimhoffiaarchimedis (Strasser, 1965) is redescribed based on several specimens collected at a number of sites close to the type locality on Sicily, Italy. Scanning electron microscopy is used to illustrate several unusual morphological characters for a member of the Glomerida for the first time. A fragment of the mitochondrial COI gene (668bp) is sequenced for the first time in Eupeyerimhoffia to provide a species-specific barcode and to gain first insights into the genetic distances between the genera in the widespread family Protoglomeridae. The novel sequences are compared to representatives of all other genera of the family: Protoglomerisvasconica (Brölemann, 1897) from northern Spain, the dwarfed Glomerellinalaurae Silvestri, 1908 from Italy and Glomeroidesprimus (Silvestri, 1929) from western North America. The addition of COI sequences from the two other families of the Glomerida renders the family Protoglomeridae paraphyletic with Glomeroidesprimus being more closely related to Glomeridellaminima (Latzel, 1884) than to the other genera in the family. The large genetic distances (13.2-16.8%) between Eupeyerimhoffia and the other genera in the order, as well as its unusual morphological characters, including unique morphological adaptations to roll into a ball, are probably an indication of the old age of the group.

As part of the German Barcode of Life (GBOL) Myriapoda program, which aims to sequence the COI barcoding fragment for 2000 specimens of Germany's 200 myriapod species in the near future, 44 sequences of the centipede order Geophilomorpha are analyzed. The analyses are limited to the genera Geophilus Leach, 1814 and Stenotaenia Koch, 1847 and include a total of six species. A special focus is Stenotaenia, of which 19 specimens from southern, western and eastern Germany could be successfully sequenced. The Stenotaenia data shows the presence of three to four vastly different (13.7-16.7% p-distance) lineages of the genus in Germany. At least two of the three lineages show a wide distribution across Germany, only the lineage including topotypes of Stenotaenialinearis shows a more restricted distribution in southern Germany. In a maximum likelihood phylogenetic analysis the Italian species Stenotaenia 'sorrentina' (Attems, 1903) groups with the different German Stenotaenialinearis clades. The strongly different Stenotaenialinearis lineages within Germany, independent of geography, are a strong hint for the presence of additional, cryptic Stenotaenia species in Germany.

Three populations of the pill millipede genus Trachysphaera Heller 1858 are present in Great Britain, one on the Isle of Wight, one in South Wales and one in mid-Wales. To identify and characterize the British Trachysphaera populations, the intraspecific and interspecific variation of the populations in South Wales and on the Isle of Wight were studied and evaluated in a first integrative study of members of Trachysphaera, utilizing barcoding and SEM. DNA was extracted from 28 British Trachysphaera and 10 French T. lobata (Ribaut 1954) specimens, one each of French T. cf. drescoi (Conde and Demange 1961) and T. pyrenaica (Ribaut 1908), and one of Spanish T. cf. rousseti (Demange 1959); the barcoding fragment of the COI gene was amplified and their genetic intra- and interpopulation distances compared with one another using two Italian T. spp. and one Croatian T. schmidti Heller 1858 specimens as near outgroups. To compare the genetic distances with the morphological characters, 15 characters of a total of 13 British Trachysphaera, together with two specimens of T.pyrenaica, two T. cf. drescoi and one of T. cf. rousseti were imaged, using the same individuals utilized for DNA extraction. Albeit both British populations are genetically distant, they are closely related (1.9-2.5% p-distance) to French T.lobata, corroborating results of earlier studies. Between different Trachysphaera species, genetic distance was high (16.7-18.8%). The morphological study showed the non-reliability of key taxonomic characters in Trachysphaera, with genetically identical individuals exhibiting morphological variation, especially on the telopods. The only observed morphological characters constant within and different between species were the number of rows of sclerotized bacilli on the tergites, as well as the shape of the male and female anal shield. Both, barcoding and the morphological study identify the British Trachysphaera populations as T. lobata.

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.

The thermophilous giant centipede Scolopendra cingulata is a voracious terrestrial predator, which uses its modified first leg pair and potent venom to capture prey. The highly variable species is the most common of the genus in Europe, occurring from Portugal in the west to Iran in the east. The northernmost occurrences are in Hungary and Romania, where it abides in small isolated fringe populations. We report the rediscovery of an isolated Austrian population of Scolopendra cingulata with the first explicit specimen records for more than 80 years and provide insights into the evolutionary history of the northernmost populations utilizing fragments of two mitochondrial genes, COI and 16S, comprising 1,155 base pairs. We test the previously proposed hypothesis of a speciation by distance scenario, which argued for a simple range expansion of the species from the southeast, via Romania, Hungary and finally to Austria, based on a comprehensive taxon sampling from seven countries, including the first European mainland samples. We argue that more complex patterns must have shaped the current distribution of S. cingulata and that the Austrian population should be viewed as an important biogeographical relict in a possible microrefugium. The unique haplotype of the Austrian population could constitute an important part of the species genetic diversity and we hope that this discovery will initiate protective measures not only for S. cingulata, but also for its habitat, since microrefugia are likely to host further rare thermophilous species. Furthermore, we take advantage of the unprecedented sampling to provide the first basic insights into the suitability of the COI fragment as a species identifying barcode within the centipede genus Scolopendra.

The redescription of the lectotype of Zephronia larvalis Butler, 1878, from the Torres Strait islands between Australia and Papua-New Guinea, shows that it does not represent a member of the SE Asian Zephroniidae, but is a species of the Australian genus Cyliosoma of the Cyliosomatidae, C. larvalis new combination. The syntypes of Cyliosoma albertisii (Silvestri, 1895; Cyliosomatidae), Australia's northern-most Sphaerotheriida species described from Somerset (close to the Torres Strait islands), were restudied, and a lectotype was selected. C. albertisii is discovered to be a junior synonym of C. larvalis (Butler, 1878). C. larvalis, originally described as Zephronia larvalis, clearly belongs to the genus Cyliosoma, but displays some characters, such as a high number (25-30) of apical cones on the antennae, and the reduction of the spine-like projection of the stigmatic plates, that are unique in the genus and family. This synonymy confirms the Torres Strait giant pill-millipede fauna to be an Australian element, and not the first representative of a still undiscovered Papua-New Guinean fauna.

The Malagasy giant pill-millipede genus Sphaeromimus de Saussure & Zehntner, 1902 is revised. Seven new species, S. titanus sp. n., S. vatovavy sp. n., S. lavasoa sp. n., S. andohahela sp. n., S. ivohibe sp. n., S. saintelucei sp. n., and S. andrahomana sp. n. were discovered, in one case with the help of sequence data, in the rainforests of southeastern Madagascar. The species are described using light- and scanning electron microscopy. A key to all 10 species of the genus is presented. All but one (S. andohahela) of the newly discovered species are microendemics each occurring in isolated forest fragments. The mitochondrial COI barcoding gene was amplified and sequenced for 18 Sphaeromimus specimens, and a dataset containing COI sequences of 28 specimens representing all Sphaeromimus species (except S. vatovavy) was analyzed. All species are genetically monophyletic. Interspecific uncorrected genetic distances were moderate (4-10%) to high (18-25%), whereas intraspecific variation is low (0-3.5%). Sequence data allowed the correct identification of three colour morphs of S. musicus, as well as the identity of a cave specimen, which although aberrant in its morphology and colouration, was genetically identical to the holotype of S. andrahoma.

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 external and internal anatomy of millipedes (Diplopoda) is poorly known compared to some of the other myriapod and arthropod groups. Due to both language barriers, which hindered the assessment of the character-rich older literature, and non-phylogenetic thinking, our knowledge of morphological characters useful for phylogenetic work diminished over the last decades. Here, a new character matrix with 64 characters, mainly derived from old literature data, is used to reconstruct a phylogeny of Diplopoda. As a tool to further our knowledge about the morphology of the different millipede orders, we show how micro-computer tomography (μCT) can be used to assess and illustrate specific parts of the Platydesmida. With the advent of μCT it is now possible to analyse many taxa and characters in a comparatively short time. A focus is put on potential phylogenetically useful characters. Our results support a Verhoeffian classification of the Diplopoda: Polyxenida + Chilognatha. Pentazonia are the sistergroup to the Helminthomorpha. Colobognatha form the sistergroup to Eugnatha, the latter split into monophyletic Juliformia and Polydesmida + Nematophora., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

We report the discovery of a ventral plate in the basal and little-known chilognath millipede order Glomeridesmida. This ventral plate, interpreted here as a 'true sternite', is clearly separate from both the coxa and the more lateral stigma-carrying plates commonly referred to as 'diplopod sternites'. Therefore, the lateral, stigma-carrying plates of the Diplopoda, previously referred to as sternites, are not sternal elements, but subcoxal elements associated with the limb base. This discovery changes the nomenclature used for the ventral plates in Diplopoda, with the formerly named 'sternite' better referred to as 'stigma-carrying plate'. In helminthomorph Diplopoda, the stigma-carrying plates are apparently secondarily fused with the sternite. The main argument for the independent evolution of tracheae in insects and myriapods, the different location of their respective spiracles, no longer holds true. In all Myriapoda and Hexapoda the spiracles associated with subcoxal elements are located lateral to the limb base. This discovery shows that the arguments for an independent origin of tracheae in insects and myriapods are not uncontestable., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

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.

To elucidate the speciation mechanisms prevalent within hotspots of biodiversity, and the evolutionary processes behind the rise of their species-rich and endemic biota, we investigated the phylogeny of the giant fire-millipede genus Aphistogoniulus Silvestri, 1897, a Malagasy endemic. This study is the first comprehensive (molecular and morphological) phylogenetic study focusing on millipede (class Diplopoda) speciation on Madagascar. The morphological analysis is based on 35 morphological characters and incorporates ten described as well as two newly described species (A. rubrodorsalisn. sp. and A. jeekelin. sp.) of Aphistogoniulus. The molecular analysis is based on both mitochondrial (COI and 16S), and nuclear genes (complete 18S rDNA), together comprised of 3031 base pairs, which were successfully sequenced for 31 individual specimens and eight species of Aphistogoniulus. In addition to the null-model (speciation by distance), two diversification models, mountain refugia and ecotone shift, were discovered to play a role in the speciation of soil arthropods on Madagascar. Mountain refugia were important in the speciation of the A. cowani clade, with three species occurring in the Andringitra and Ranomafana Mountains in the southeast (A. cowani), the Ambohijanahary and Ambohitantely Mountains in the mid-west (A. sanguineus), and the Marojejy Mountain in the northeast (A. rubrodorsalisn. sp.). An ecotone shift from the eastern rainforest to the unique subarid spiny forest of Mahavelo was discovered in the A. vampyrus-A. aridus species-pair. In the monophyletic A. diabolicus clade, evidence for divergent evolution of sexual morphology was detected: species with greatly enlarged gonopods are sister-taxa to species with normal sized gonopods. Among the large-bodied Spirobolida genera of Madagascar, Colossobolus and Sanguinobolus were found to be close sister-genera to Aphistogoniulus. Forest destruction has caused forest corridors between populations to disappear, which might limit the possible resolution of biogeographic analyses on Madagascar.