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The millipedes of Yintiaoling National Natural Reserve, Southwest China, were investigated. In total, thirteen species from five orders (Sphaerotheriida, Spirobolida, Callipodida, Chordeumatida and Polydesmida) and eight families were discovered. Among them, six species are new to science: Zephronia linkouzi sp. nov., Paracortina inflata sp. nov., Orthomorpha laminata sp. nov., Polylobosoma corollifera sp. nov., Epanerchodus wuxi sp. nov. and Riukiaria spina sp. nov. As usual, the polydesmidan family Paradoxosomatidae is the most diverse group, containing five species. All new species are described and illustrated, and photographs of the habitus and the gonopods of all species are also provided.

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

A new species of giant pill-millipede is described and illustrated from Guizhou Provinces, southern China: Sphaerobelum huzhengkuni n. sp. This is the first record of the genus Sphaerobelum in Chin...

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 genusZoosphaeriumPocock, 1895 which are described here asZoosphaerium mangabeWesener,sp. nov.,Z. bartolozziiAnilkumar & Wesener,sp. nov., andZ. taitiiAnilkumar & Wesener,sp. nov., all belonging to theZ. coquerelianumspecies 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 granulatusGervais, 1847, andChondromorpha xanthotricha(Attems, 1898) are recorded for the first time from Madagascar. New locality data is provided forZoosphaerium neptunus(Butler, 1872),Z. villosumWesener & Sierwald, 2005,Z. blandum(de Saussure & Zehntner, 1897),Sphaeromimus musicus(de Saussure & Zehntner, 1897),Rhinotus purpureus(Pocock, 1894),Hylekobolus andasibensisWesener, 2009,Aphistogoniulus infernalisWesener, 2009,Ostinobolus rufusWesener, 2009,Ostinobolus subterraneusWesener, 2009,Dactylobolus bivirgatus(Karsch, 1881), andEumekius antimena(de Saussure & Zehntner, 1901).

Three new species of the giant pill-millipede family Zephroniidae are described from southern Vietnam: Sphaerobelum pumatensesp. nov., Sphaeropoeus honbaensissp. nov. and Sphaeropoeus bidoupensissp. 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.

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.

Seven new species of the giant pill-millipede genus Zoosphaerium Pocock, 1895 are described from Madagascar: Z. nigrum sp. nov., Z. silens sp. nov., Z. ambatovaky sp. nov., Z. beanka sp. nov., Z. voahangy sp. nov., Z. masoala sp. nov. and Z. spinopiligerum sp. nov. All species are described based on drawings and scanning electron microscopy, while genetic barcoding of the COI gene was successful for six of the seven new species. Additional COI barcode information is provided for the first time for Z. album Wesener, 2009 and Z. libidinosum (de Saussure & Zehntner, 1897). Zoosphaerium nigrum sp. nov. and Z. silens sp. nov. belong to the Z. libidinosum species-group, Z. ambatovaky sp. nov. to the Z. coquerelianum species-group, Z. beanka sp. nov., Z. voahangy sp. nov. and Z. masoala sp. nov. to the Z. platylabum species-group and Z. spinopiligerum sp. nov. to the Z. piligerum species-group. Updated identification keys are provided for each species-group. Two western dry forest species, Z. silens sp. nov. and Z. voahangy sp. nov. are recorded from two localities, while the other five species are currently only known from their type localities. Of special conservation concern might be Z. ambatovaky sp. nov. from the lowland rainforest fragment of Ambatovaky, a nowadays isolated lowland rainforest, rapidly shrinking due to slash and burn agriculture. In addition to the new species, new locality data is provided for 11 species and numerous unidentifiable species of Zoosphaerium: Z. neptunus (Butler, 1872), Z. platylabum (de Saussure & Zehntner, 1902) and Z. piligerum (de Saussure & Zehntner, 1897) from the central eastern montane forests, as well as Z. ambrense Wesener, 2009, Z. aureum Wesener, 2009, Z. libidinosum, Z. corystoides Wesener, 2009, Z. discolor Wesener, 2009, Z. smaragdinum Wesener, 2009, Z. villosum Wesener & Sierwald, 2005 and Microsphaerotherium anjozorobe Wesener, 2009.

Two new species of giant pill-millipedes, Zephronia viridisoma Rosenmejer & Wesener sp. nov. and Sphaerobelum aesculus Rosenmejer & Wesener sp. nov., are described based on museum samples from southern Thailand. Zephronia viridisoma sp. nov. comes from Khao Lak, while the type locality of S. aesculus sp. nov. is on Phuket Island. Both species are described integratively, combining light microscopy, scanning electron microscopy, multi-layer photography, micro-CT scans and genetic barcoding. Genetic barcoding was successfully conducted for holotypes of both new species, which could be added to a dataset of all published sequences of the family Zephroniidae, including all described species from Thailand, Laos and Cambodia up to 2020. Genetic barcoding of the COI gene revealed another female of S. aesculus sp. nov., 160 km east of the type locality. Both new species are genetically distant from all other Zephroniidae from Thailand and surrounding countries, showing uncorrected p-distances of 16.8–23.1%. A virtual cybertype of a paratype of Z. viridisoma sp. nov. was created and made publically accessible.

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.

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.

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 endemic giant pill-millipedes belonging to the genus Arthrosphaera has a long history of occurrence and distribution in the Indian Subcontinent. According to the earlier and recent records, up to 40 species of Arthrosphaera are confined to southern India and Sri Lanka. Being detritus feeders, they are sensitive to narrow range of abiotic factors and their population is dwindling due to severe human interference especially landscape modification or modern forestry/agricultural practices. There is an urgent need to update the status, occurrence and distribution in distinct ecological conditions of pill-millipedes in southern India as they are important component in organic matter decomposition as well as nutrient turn over in the forests or plantations. Based on recent surveys, the present study attempts to update occurrence, distribution and morphological description (light and scanning electron microscopy) with a diagnostic key for identification of 11 species of pill-millipedes occurring in the Western Ghats of India with commentary on their distribution pattern in different ecological conditions and impact of edaphic factors.

La Camelia bei Angelopolis. Unspecified number of ♂ and ♀. The MHNG collection contains three specimens in alcohol. The specimens each have a pin running the length of the body and are in a glass vial (MHNG- ARTO-14357). A smaller vial (MHNG-ARTO-14358) contains gonopods. The identification labels in the jar both have “ Colombie coll. Fuhrmann ” written on them and there is a printed “Cotype” label, indicating that the specimens are syntypes. There are also two syntypes (one ♂ and one ♀ referred to as “ Co-Typus ” in the NMB catalogue) in the NMB (inventory number NMB-DIPL- 00227a). Trichomorpha annulipes Carl, 1914, Chelodesmidae, Published as part of Hollier, John, Stöckli, Eduard, Wesener, Thomas, Mesibov, Robert, Decker, Peter & Sierwald, Petra, 2020, An annotated list of the millipede (Diplopoda) species described by Johann Carl, pp. 183-240 in Revue suisse de Zoologie 127 (1) on page 187, DOI: 10.35929/RSZ.0015, http://zenodo.org/record/5743454

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.

Along with plant sources, arthropods (such as millipedes) are also known for their antioxidant activities that can help combat human ailments. Therefore, methanolic extracts of males and females of two endemic giant pill-millipedes of the Western Ghats (Arthrosphaera fumosaPocock andArthrosphaera magnaAttems) were evaluated for antioxidant potential. The bioactive principles and activities assessed were total phenolics, total antioxidant activity, 2,2-diphenyl-1-picrylhydrazyl or DPPH radical scavenging, ferrous ion chelation, reducing power and hydrogen peroxidase scavenging. Significant differences were shown between the two species, the sexes and the sexes within millipede species in total antioxidant activity, DPPH radical scavenging and ferrous ion chelation. Total phenolics and EC50of DPPH radical scavenging activity differed significantly only between species and the sexes. This study clearly demonstrated that the antioxidant capacity of the methanolic extract ofArthrosphaeraspp. is comparable to the ascorbic acid standard and possesses potential scavenging ability against reactive oxygen species generated during oxidative stress.

Certain groups of invertebrates are becoming mainstreamed in conservation activities in KwaZulu-Natal, particularly groups that have many species endemic to the province and that are confined to small distribution ranges. As a result of this there is a need to assign common (English) names where these are unavailable. A list of scientific names and standardised common names is presented for species of millipede occurring in KwaZulu-Natal.

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.

Mating behaviour of four species of pill-millipedes under genus Arthrosphaera Pocock (Arthrosphaera dalyi Pocock and Arthrosphaera disticta Pocock, Arthrosphaera fumosa Pocock and Arthrosphaera magna Attems) endemic to the Western Ghats of Southern India was analyzed in mesocosms. Stridulation is a classical communication signal in males as well as females for mate selection. Conglobation (or volvation) is a mechanism of defence to protect from disturbance or avoid predation. If male touches female or vice versa they conglobate. To avoid disparity among individuals of the same species, volvating pill-millipedes evolved stridulation behaviour for communication. The male broadcasts appropriate signal to female through stridulation to advertise its interest in mating. The females test the male’s fitness by conglobation and suitable male uncoils the partner through stridulation signals. Male with its pygidium successfully uncoils the female and attains suitable orientation for courtship. Male pairs with female ventro–ventro contact in opposite direction to deposit sperm into the vulva of female. The duration of mating varies from species to species and usually a lapse from 3 to 30 min. Vibration generated by stridulation is species-specific and its perception mechanism in pill-millipedes is yet to be clearly understood. Present study emphasized the structure of stridulatory organs, mechanism of stridulation and pattern of mating behaviour in four species of pill-millipedes.

The species-rich giant pill-millipedes (Sphaerotheriida) often represent a microendemic component of Madagascar’s mega-invertebrate fauna. Of the chirping genus Sphaeromimus de Saussure & Zehntner, 1902, ten species have been described. Here, we describe two new species of Sphaeromimus integratively, combining light microscopy, scanning electron microscopy, DNA barcoding and micro-CT technology for the first time in a taxonomic description of a giant pill-millipede. S. kalambatritra sp. nov. and S. midongy sp. nov. are the first giant pill-millipedes collected and described from the mountainous rainforests of Kalambatritra and Midongy. Both species show island gigantism compared to their congeners. Our analysis of the mitochondrial COI gene shows that the two species are related to one another with a moderate genetic distance (9.4%), while they are more closely related to an undetermined specimen from the forest of Vevembe (6.3% and 8.4%). They stand in a basal position with S. ivohibe Wesener, 2014 and S. musicus (de Saussure & Zehntner, 1897). The four aforementioned species share a high number of stridulation ribs on the male harp. Our micro-CT analysis provides a look into the head of S. kalambatritra sp. nov. and shows that non-destructive CT methods are a useful tool for studying the inner morphology of giant pill-millipedes.

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

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 >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.

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.

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

Five new species of the endemic giant pill-millipede genus Zoosphaerium from Madagascar are described: Z. muscorum sp. n., Z. bambusoides sp. n., Z. tigrioculatum sp. n., Z. darthvaderi sp. n., and Z. heleios sp. n. The first three species fit into the Z. coquerelianum species-group, where Z. tigrioculatum seems to be closely related to Z. isalo Wesener, 2009 and Z. bilobum Wesener, 2009. Z. tigrioculatum is also the giant pill-millipede species currently known from the highest elevation, up 2000 m on Mt Andringitra. Z. muscorum and Z. darthvaderi were both collected in mossy forest at over 1000 m (albeit at distant localities), obviously a previously undersampled ecosystem. Z. darthvaderi and Z. heleios both possess very unusual characters, not permitting their placement in any existing species-group, putting them in an isolated position. The females of Z. xerophilum Wesener, 2009 and Z. pseudoplatylabum Wesener, 2009 are described for the first time, both from samples collected close to their type localities. The vulva and washboard of Z. pseudoplatylabum fit very well into the Z. platylabum species-group. Additional locality and specimen information is given for nine species of Zoosphaerium: Z. neptunus (Butler, 1872); Z. villosum Wesener & Sierwald, 2005; Z. amabile Wesener, 2009; Z. tampolo Wesener, 2009; Z. smaragdinum Wesener, 2009; Z. viridissimum, Wesener, 2009; Z. pseudopiligerum Wesener, 2009; and Z. analavelona Wesener, 2009.

Various ants can link their bodies in chains or other structures, a striking example of cooperation. Chain behavior in Leptogenys sp. functions for the collective transport of large millipedes. This undescribed Leptogenys species is a swarm raider with a small range of prey: millipedes belonging to four orders (Polydesmida, Spirostreptida, Spirobolida, Sphaerotheriida), and occasional earthworms. Small prey were carried individually or dragged by a few ants, while chains made it possible to move millipedes weighing up to 16.4 g. Chains (either linear or branched) changed according to obstacles along the way to the nests. Between 2 and 52 workers were observed to drag single prey items, with only a few ants directly grasping the prey. One 15-cm-long millipede was captured rodeo-style after being encircled by 25–30 ants. As it uncurled from a defensive coil, the ants held back except one that tried to sting between its legs. The millipede started thrashing about which caused many ants to attempt stinging. While four other ponerine genera hunt millipedes solitarily, chains have never been reported.

The millipede Anaulaciulus koreanus (Verhoeff, 1937), belonging to the family Julidae, is an endemic species of the Korean fauna. In this study, we sequence and annotate the mitochondrial genome of A. koreanus . The complete mitochondrial genome of this species is 14,916 bp in length and contains 13 protein-coding genes, 22 transfer RNA genes, two ribosomal RNA genes (16S and 12S rRNA), and a large non-coding region. The genome has a very high A+T content (71.1%), less than of the species Brachycybe lecontii Wood, 1864 (order Platydesmida; 76.6%) and Sphaerotheriidae sp. (order Sphaerotheriida; 71.2%). In comparison with the mitochondrial gene arrangement of eight other millipede species, the whole mitochondrial gene arrangement of A. koreanus is most similar to the nemasomatid species, Antrokoreana gracilipes Verhoeff, 1938, but differs from those of the other diplopod orders. The absence of tRNA Cys between the ND2 and COI regions is unique to the order Polydesmida, whereas the translocation of tRNA Tyr to between ND2 and COI is exclusive to the Sphaerotheriida. It is also shown that the translocation of tRNA Thr between ND4L and ND1 may be a synapomorphy to support a close relationship of two orders Spirobolida and Spirostreptida.

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.

Thirty-one new species of giant pill-millipedes (order Sphaerotheriida) endemic to Madagascar are described: Zoosphaerium haackeri n. sp., Z. album n. sp., Z. discolor n. sp., Z. mitoho n. sp., Z. pulchellum n. sp., Z. endemicum n. sp., Z. xerophilum n. sp., Z. aureum n. sp., Z. pseudopriapus n. sp., Z. pseudoplatylabum n. sp., Z. solitarium n. sp., Z. tsingy n. sp., Z. corystoides n. sp., Z. broelemanni n. sp., Z. amabile n. sp., Z. trichordum n. sp., Z. smaragdinum n. sp., Z. ambrense n. sp., Z. isalo n. sp., Z. bilobum n. sp., Z. tampolo n. sp., Z. ignotum n. sp., Z. tainkintana n. sp., Z. viridissimum n. sp., Z. denticulatum n. sp., Z. pseudoblandum n. sp., Z. micropiligerum n. sp., Z. pseudopiligerum n. sp., Z. analavelona n. sp. and Z. fisheri n. sp., and Microsphaerotherium anjozorobe n. sp.. Twenty-eight of the Zoosphaerium species already fit into established species-groups. A key to all 55 recognizable Malagasy Sphaerotheriida species is given. Forty-two of the 55 giant pill-millipede species are ecosystem endemics, while 26 are geographically microendemic. The number of microendemic species is disproportionately distributed among Malagasy ecosystems. All 13 species restricted to the montane rainforest as well as all five species restricted to the Eastern littoral forest are microendemic. On the other hand, only two out of seven dry forest species, one out of seven rainforest species and one out of five spiny forest species are microendemics. A special emphasis is placed on two species endemic to the unique Western rainforest site of Analavelona, which is currently not protected.

This review lists the currently known species of millepedes in Sri Lanka and discusses their current taxonomic status and distribution based on previous studies from 1865 to date. A total of 104 millipede species belonging to 44 genera, 18 families and nine orders have been recorded in Sri Lanka. Of these, 82 are known only from Sri Lanka; additionally, nine genera and one family are known only from Sri Lanka. Most of the millipede species have been recorded from two localities, namely Pundaluoya and Kandy in the central highlands of Sri Lanka. Current knowledge on the taxonomy, evolutionary relationships, distribution and conservation of the millipedes of Sri Lanka is still limited and scattered. Thus we suggest more intensive surveys to acquire comprehensive data on the millipedes of Sri Lanka.

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 redescription of the lectotype of Z ephronia 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.

At the present, the millipede fauna of Laos comprises only 34 species from 20 genera, 13 families and 7 orders. These counts certainly represent but a minor fraction of the country’s real diversity of Diplopoda even at the ordinal level, let alone at lower ones. Based on the available information from the adjacent parts of China, Thailand, Myanmar, Vietnam and/or Cambodia, the orders Polyxenida, Sphaerotheriida, Chordeumatida, Julida and Polyzoniida must occur in Laos, may be also Glomeridesmida, Siphonocryptida and Stemmiulida, but none has been recorded there yet. Moreover, even some ubiquitous “tramp” species, such as Glyphiulus granulatus (Gervais, 1847), Trigoniulus corallinus (Gervais, 1847), Desmoxytes planata (Pocock, 1895) or Oxidus gracilis (C. L. Koch, 1847), have hitherto not been found in Laos. This shows that a lot more collecting efforts, which have heretofore been rather strongly biased to caves, are required to amass a representative material of Diplopoda of Laos and make it available for study.

Cyliosoma Pocock, 1895, the oldest available genus name for Australian giant pill-millipedes, is revised with a redescription of its type species, Sphaerotherium angulatum Butler, 1878. All 16 species of Epicyliosoma Silvestri, 1917 are transferred to Cyliosoma, together with two species, Sphaerotherium fraternum Butler, 1872 and S. marginepunctatum Karsch, 1881, which are redescribed here. A new phylogenetic analysis of the Sphaerotheriida was conducted using 100 morphological characters and including two Cyliosoma species and four recently described or redescribed species of the family Zephroniidae. Most character states are illustrated for Cyliosoma, including the first SEM images of a member of the genus. Cyliosoma is neither closely related to the South African Sphaerotherium, nor to the other Australian genus, Procyliosoma, and is here placed in a new family, Cyliosomatidae. The monotypic Australian genus Cynotelopus Jeekel, 1986 is also referred to the Cyliosomatidae. The current position of the Cyliosomatidae is in a trichotomy including the South African Sphaerotheriidae and the Malagasy–Indian Arthrosphaeridae.

As a first step towards an inventory of the giant pill-millipedes in Thailand, a new species of the genus Sphaerobelum Verhoeff, 1924, S. truncatum n. sp. is described from Nan Province, northern Thailand. A determination key is presentedfor all five known Sphaerobelum species. Clear morphological differences between S. truncatum n. sp. and the other four Sphaerobelum species were found on the anterior telopods. For the first time in Sphaerobelum, the partial mitochondrialCOI gene was sequenced for S. truncatum n. sp. and compared with distance, maximum parsimony and maximum likeli-hood methods to those of species from other giant pill-millipede genera. Sphaerobelum truncatum n. sp. was found to dif-fer from all other analyzed giant pill-millipedes, including species of Zephronia Gray, 1832, by 22–30%, includingnumerous amino acid changes, supporting the separate status of Sphaerobelum among other giant pill-millipede genera.Maximum likelihood and parsimony analyses support the placement of Sphaerobelum in the Zephroniidae. Figures of allrelevant structures of Sphaerobelum truncatum n. sp. are provided to allow the use of these characters in future descriptions of species of the family Zephroniidae.

Two new species of the Malagasy sphaerotheriid genus Zoosphaerium, Z. villosum sp. nov., and Z. arborealis sp. nov., are described. Characters of a shiny black new putative species of the genus Zoosphaerium are described and illustrated. Zoosphaerium alluaudi DeSaussure & Zehntner is redescribed. The characters defining the genus Zoosphaerium are summarized: three jointed anterior telopods, first joint with a stridulation organ termed ‘harp’ composed out of 1–2 stridulation ribs. Females with stridulation ribs on the subanal plate, termed ‘washboard’. Operculum of female vulva constricted in the middle (‘subreniform’). 6 th joint of antennae of cylindrical shape. The nomenclatural history of the genus name is discussed. The intraspecific variability of many characters commonly used to delineate species within the genus Zoosphaerium and other members of the order Sphaerotheriida is analyzed in detail. Ontogenetic changes in the female vulva and the male telopods are discussed and illustrated.Zwei neue Arten der Madagassischen Gattung Zoosphaerium, Z. villosum sp. nov. und Z. arborealis sp. nov., werden beschrieben. Merkmale einer wahrscheinlich neuen, aber hier nicht benannten Art der Gattung werden beschrieben und illustriert. Die Art Zoosphaerium alluaudi DeSaussure & Zehntner, 1902 wird wiederbeschrieben. Die nomenklatorische Geschichte und die Merkmale der Gattung Zoosphaerium werden zusammengefasst. Mitglieder der Gattung zeichnen sich durch folgende Merkmale aus: Dreigliedrige anteriore Telopoden mit einem Stridulationsorgan auf dem ersten Glied, welches aus 1–2 Stridulationsrippen besteht und als Harfe bezeichnet wird. Weibchen mit mehreren Stridulationsrippen auf der Subanalplatte, „washboard“ genannt. Operculum der weiblichen Vulva am Oberrand in der Mitte eingebuchtet (subreniform). 6. Antennenglied zylindrisch geformt. Die intraspezifische Variabilität mehrerer in der Gattung Zoosphaerium häufig benutzter Artmerkmale wurde untersucht. Formenwandel der weiblichen Vulven und männlicher Telopoden während der Ontogenie werden beschrieben und abgebildet.