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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.

The first new fossil pill millipede, order Glomerida, description in more than 150 years is based on two specimens, one male and one female, found in Baltic amber. Both specimens were studied with light- and micro-CT technology, allowing a detailed reconstruction of the posterior telopods and numerous other morphological characters. Based on these characters, the specimens can be securely placed in the megadiverse, mainly Asiatic genus Hyleoglomeris Verhoeff, 1910 as a new species, Hyleoglomeris groehni n. sp. The species H. groehni n. sp. is the oldest and only known fossil member of the Glomerida that is sufficiently described to allow for placement in a family or genus. The discovery of a fossil Hyleoglomeris in Baltic amber has biogeographic implications as the genus is now speciose in Asia, with the nearest relatives found in the Caucasus and some caves in Serbia and Greece. With the discovery of a fossil European Hyleoglomeris species, the cave Hyleoglomeris species are interpreted as relicts, surviving underground as living witnesses of a different time when Hyleoglomeris was still extant in Europe. Other cave Glomerida in Europe, as well as morphologically aberrant pill millipede species endemic to the Canary Islands, should be checked for potential placement in Hyleoglomeris.

A new pill millipede, Rhopalomeris nagao sp. nov., is described from Vietnam. It is recognized by the following combination of characters: antennal tip with numerous sensory cones; telopods with short prefemoral and femoral trichosteles, with long, straight and acute tuberculiform femoral process, and with shorter lobuliform tibial process; syncoxial lobe being slightly concave medially, syncoxial horns being longer than lobe. A fragment of the cytochrome c oxidase subunit I (COI) gene is also provided for the new species. The K2P genetic distance of the COI between the new species and other Vietnamese glomeridans is from 10.7% to 16.9%. Two genera, Hyperglomeris and Hyleoglomeris, are considered to be non-monophyletic, but more data and samples will be needed for confirmation.

The Black Pill Millipede,Glomeris marginata, is the best studied millipede species and a model organism for Diplopoda.Glomeris marginatais 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 ofG. marginata. The results of the COI barcoding fragment analysis include 97G. marginata, as well as 21 specimens from seven potentially related species:G. intermediaLatzel, 1884,G. klugiiBrandt, 1833 (G. undulataC.L. Koch, 1844),G. connexaKoch, 1847,G. hexastichaBrandt, 1833,G. maerensAttems, 1927,G. annulataBrandt, 1833 andG. apuanaVerhoeff, 1911. The majority of the barcoding data was obtained through the German Barcode of Life project (GBOL). Interspecifically,G. marginatais separated from its congeners by a minimum uncorrected genetic distance of 12.9 %, confirming its monophyly. Uncorrected intraspecific distances ofG. marginataare comparable to those of other widespreadGlomerisspecies, varying between 0–4.7%, with the largest genetic distances (>2.5 %) found at the Mediterranean coast. 97 sampled specimens ofG. marginatayielded 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 inG. 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.

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 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.

This paper reports on an abnormally developed embryo (ADE) of the common pill millipede Glomeris marginata. This ADE represents a modified case of Duplicitas posterior, in which two posterior ends are present, but only one anterior end. While the major posterior germ band of the embryo appears almost normally developed, the minor posterior germ band is heavily malformed, has no clear correlation to the single head, little or no ventral tissue, and a minute amount of yolk. The anterior end of the minor germ band is fused to the ventral side of the major germ band between the first and second trunk segment. At least one appendage of the second trunk segment appears to be shared by the two germ bands. Morphology and position of the minor germ band suggest that the ADE may be the result of an incorrectly established single cumulus [the later posterior segment addition zone (SAZ)]. This differs from earlier reports on D. posterior type ADEs in G. marginata that are likely the result of the early formation of two separate cumuli.

A new genus and species of the pill millipede order Glomerida, Nearctomeris inexpectata n. sp., n. gen., is describedfrom the Eastern United States. The description of Nearctomeris brings the total number of American Glomerida generato three. In two of three known localitites, Nearctomeris co-occurs with species of the genus Onomeris Cook, 1896. Onomeris and Nearctomeris have almost an identical size (4–5.5 mm) and dark colour, but can be readily distingusihedby numerous non-sexual and sexual characters. Sexual characters of the male telopod also place Nearctomeris into thefamily Glomeridae, while the third American pill millipede genus Glomeroides Chamberlin, 1922 belongs to theProtoglomeridae. A key to all American genera of Glomerida is presented. For the first time in the order Glomerida, thepartial cytochrome c oxidase I mitochondrial gene was analyzed for the three American genera of the Glomerida, Glomeroides, Onomeris, and Nearctomeris together with European genera currently placed in different families, Glomeridella and Trachysphaera, as outgroups. Distance, maximum parsimony and maximum likelihood methods wereemployed. Maximum parsimony and likelihood analyses did not lead to a well-resolved phylogeny, but found a weaklysupported sister-group relationship between Nearctomeris and Onomeris. All five analyzed pill millipede genera differgreatly in their uncorrected basepairs (15–22%), and moderately in their amino acids (5–10%). As in Onomeris, the closestrelative of Nearctomeris can probably be found among the Asian genera Hyleoglomeris Verhoeff, 1910 or Hyperglomeris Silvestri, 1917. In America, more attention should be given to the search for other small-bodied, cryptic, rare Glomerida.

Seven species of the basically warm temperate to tropical Eurasian genus Hyleoglomeris are currently known from Thailand, including three new ones: H. hongkhraiensis sp. n. and H. aurea sp. n. from Chiang Mai Province, and H. cavicola sp. n. from Sukhothai Province. A new distribution map and a key to all Hyleoglomeris species presently known to occur in Thailand are given.

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 >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 >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 Rothenbuhler, 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.

In some arthropods there is a discrepancy in the number of dorsal tergites compared to the number of ventral sternites and leg pairs. The posterior tergites of the Diplopoda (millipedes) each cover two sternites and two pairs of legs. This segment arrangement is called diplosegmentation. The molecular nature of diplosegmentation is still unknown. There are even conflicting theories on the way the tergites and sternites/leg pairs should be correlated to each other. The different theories are based either on embryological analyses or on studies of the adult morphology and turned out to be not compatible with each other. We have previously used the expression patterns of segmentation genes in the pill millipede Glomeris marginata (Myriapoda: Diplopoda) to study millipede segmentation. Here we review the existing models on the alignment of tergites and leg pairs in millipedes with special emphasis on the implications the gene expression data have on the debate of tergite and leg pair assignment in millipedes. The remarkable outcome of the gene expression analysis was that (1) there is no coupling of dorsal and ventral segmentation and, importantly, that (2) the boundaries delimiting the tergites do neither correlate to the embryonic boundaries of the dorsal embryonic segments nor to the boundaries of the ventral embryonic segments. Using these new insights, we critically reinvestigated the correlation of tergites, sternites, and leg pairs in millipedes. Our model, which takes into account that the tergite boundaries are different from the dorsal embryonic segment boundaries, provides a solution of the problem of tergite to sternite/leg pair correlation in basal milipedes with non-fused exoskeletal elements and also has implications for derived species with exoskeletal rings. Moreover, lack of coupling of dorsal and ventral segmentation may also explain the discrepancy in numbers of dorsal tergites and ventral leg pairs seen in some other arthropods.

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.