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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.

Two new pill millipede species, namely Hyleoglomeris hoanglien sp. nov. and H. fanxipan sp. nov., are described from high elevations (approximately 2,800–2,900 m) of the Hoang Lien National Park, n...

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.

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 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.

Rensch’s rule predict the negative associations between sexual size dimorphism (SSD) and body sizes for relatively larger females. This prediction was tested for forest and savanna diplopods using a geometric morphometric approach using calculations of length and width to derive shape volume based on the mathematical formulae for estimated cylindrical worm-like millipede size (l.π.r2) and spherical pill millipede size (4/3.π.r3): (i) Centrobolus (were collected in February 1996, South Africa) SSD was 0.63-2.89 (1.52±0.35; 267) (ii) Sphaerotherium (extracted from literature) SSD was 1.49-5.36 (2.96±1.40; n≥7); (iii) savanna (Calostreptus, Doratogonus, Odontopyge and Spinotarsus were collected in February 1989, Zimbabwe) and forest helminthomorphs (collected in February 1996, South Africa) SSD was 0.88-1.62 (1.26±0.23; 1233). Interspecific variation regressed was SSD (0.63-5.36) on body sizes (n=1273) with no significant negative correlations rejecting Rensch’s rule. Eco-morphological patterns were discussed.

Rensch’s rule predict the negative associations between sexual size dimorphism (SSD) and body sizes for relatively larger females. This prediction was tested for forest and savanna diplopods using a geometric morphometric approach using calculations of length and width to derive shape volume based on the mathematical formulae for estimated cylindrical worm-like millipede size (l.π.r2) and spherical pill millipede size (4/3.π.r3): (i) Centrobolus (were collected in February 1996, South Africa) SSD was 0.63-2.89 (1.52±0.35; 267) (ii) Sphaerotherium (extracted from literature) SSD was 1.49-5.36 (2.96±1.40; n≥7); (iii) savanna (Calostreptus, Doratogonus, Odontopyge and Spinotarsus were collected in February 1989, Zimbabwe) and forest helminthomorphs (collected in February 1996, South Africa) SSD was 0.88-1.62 (1.26±0.23; 1233). Interspecific variation regressed was SSD (0.63-5.36) on body sizes (n=1273) with no significant negative correlations rejecting Rensch’s rule. Eco-morphological patterns were discussed.

Sexual size dimorphism (SSD) was investigated in the millipede genus Sphaerotherium. Width was extracted from Attems (1928) monograph and used to compare interspecific variation in mean volumes using a geometric morphometric approach. Based on the formula for a sphere (4/3. π. r3), volume was calculated in seven species and an allometric coefficient of 0.7 found. The allometric equation generated for the genus was ŷ = 0.00251X + 57211.22445. Correlation between SSD and body sizes (R = 0.749728; P = 0.00202; n=7) does now reject Rensch’s rule in Sphaerotherium.

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.

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 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.

SUMMARY All trunk segments in the pill millipede Glomeris marginata (Myriapoda: Diplopoda) are initially patterned genetically, (as visualized by the embryonic expression pattern of the even-skipped gene) and formed morphologically, (as visualized by 4-6-diamidin-2-phenylindol stained embryos) in a single segmental period. In addition, formation of every nascent trunk segment concerns ventral as well as dorsal segmental units. Only after the formation of the nascent posterior trunk segments, the dorsal segmental units of two adjacent segments fuse to form a single dorsal segmental unit that subsequently covers two ventral leg-bearing segmental units. The formation of a diplosegmental unit, or in short a diplosegment, is thus the result of dorsal fusion of embryonic tissue and not the result of any splitting-process or fusion of dorsal tergites. The new data also argue against heterochrony as a primary causative factor for the formation of the diplosegments during the formation of dorsal versus ventral segmental units. Furthermore, no evidence was found supporting the hypothesis that anterior trunk segments in diplopods represent degenerate diplosegments. Two possible scenarios arise from the ontogenetic data presented here, whether this represents an ancestral feature of the diplopods, or alternatively if they represent an isolated case only found in Glomeris (and close relatives). If the former is the case, my work may provide an impressive example of Haeckel's recapitulation theory.

Specimens of the North American glomerid millipede Onomeris sinuata (Loomis, 1943) were collected at the type locality in Alabama, USA, and maintained briefly in the laboratory in native leaf litter. The millipedes could not be induced to produce defensive secretions by rough handling, tapping, squeezing or leg-pinching. Four specimens were extracted in methanol and the extract analyzed using gas chroma-tography/mass spectroscopy. The analysis revealed the alkaloids glomerin and homoglomerin, previously reported as components in the defensive secretion of the European glomerid Glomeris marginata (Villers, 1789). This report is only the second for the occurrence of quinazolinone alkaloids in animals. Keywords Onomeris sinuata , Glomeris marginata , quinazolinone alkaloids, Alabama, USA Introduction Nearly simultaneously in 1966, Schildknecht et al. (1966)and Meinwald et al. (1966) announced the discovery of two quinazolinone alkaloids in the defensive secre-tion of the European millipede Glomeris marginata

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.

Spinous oniscomorph millipedes are rare faunal components of the upper Palaeozoic (Shear, 1997). Amynilyspes (type species A. wortheni Scudder, 1882; OD) is an Upper Carboniferous spinous oniscomorph (pill millipede) which was first described from the Middle Pennsylvanian (Westphalian D equivalent) Fossil-Lagerstatte of Mazon Creek. Later Fritsch (1899) described two species ( A. typicus and A. crescens ) from the Wesphalian D Gaskohle of Nýřany in Bohemia. More recently A. typicus was recorded from the Stephanian B of the Saarland (Forster, 1973), then from the Stephanian B of the Blanzy-Montceau-les-Mines Basin (Langiaux and Sotty, 1977; Poplin, 1994). Although myriapods were already known to occur in the Montceau-les-Mines Lagerstatte (Langiaux and Sotty, 1976; Rolfe et al., 1982; Poplin and Heyler, 1994) the presence of Amynilyspes at Montceau-les-Mines was first published in a regional publication (“La Physiophile” Langiaux and Sotty, 1976, 1977), and it has passed almost unnoticed. Representatives of the genus Amynilyspes are a minor faunal component of Upper Carboniferous faunal assemblages. Hannibal and Feldmann (1981) published a revision of the oniscomorph millipedes from the Carboniferous of North America, including the type specimen of Amynilyspes wortheni Scudder, 1882 and newly collected specimens. Specimens of the genus Amynilyspes subsequently have been illustrated and discussed in various works (Hannibal, 1984, 1997; Shear, 1997). At Mazon Creek Hannibal and Feldmann (1981) listed 10 specimens of A. wortheni , plus an undetermined number of specimens in private collections. The percentages of the whole millipede fauna (excluding arthropleurids) are 0.1 percent in the Braidwood (nonmarine, freshwater) biofacies and 0.014 percent in the Essex (marine) environment (Baird and Anderson, 1997). A survey of fossil millipedes in the collections of the Field Museum of Natural History by one of us (JTH) in 1997 showed that oniscomorphs (mainly Amynilyspes ) comprise about 11 percent (24 …

The complete mitochondrial genome (mitogenome) of the millipede Sphaerotheriidae sp. has been studied. The genome is 14,970 bp long and contains the typical complement of 13 protein-coding genes, 22 transfer RNA genes, and 2 ribosomal RNA genes. Gene order in Sphaerotheriidae sp. mitogenome is assumed to represent the myriapod ground pattern, which is shared by myriapod-chelicerate clade.

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.

A new genus of Laboulbeniales, Triaino- myces, is described and illustrated based on a single species, T. hollowayanus found parasitizing females of the pill-millipede Procyliosoma tuberculatum in New Zealand. Structure and development of the thallus of T hollowayanus is described and distinctive features are discussed. The new genus is placed in the subtri- be Peyritschiellinae of the Peyritschielleae although perithecial ontogeny is similar to that observed in members of the Laboulbenioideae.

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.

Abnormally developing embryos (ADEs) of the common pill millipede Glomeris marginata have been investigated by means of nuclear staining and mRNA in situ hybridization. It showed that all ADEs represent cases of Duplicitas posterior, which means that the posterior body pole is duplicated. The severity of the duplication ranges from duplicated posterior trunk segments in one specimen to an almost completely duplicated specimen that only shares the very anterior head region. Remarkably, none of the encountered ADEs represents a case of Duplicitas anterior (duplicated anterior pole) or a case of Duplicitas cruciata (cruciate duplication with two anterior and two posterior poles). This observation is discussed in the light of earlier reports on G. marginata ADEs that claim to have found these abnormalities. The lack of any other axial abnormality aside from D. posterior implies that early axis determination in G. marginata, and possibly myriapods in general, underlies the developmental mechanisms that prevent the formation of any other type of axial duplication. It is proposed that the formation of D. posterior-type embryos could be caused by the formation of two instead of only one posterior cumulus early during development.

The body of arthropods is subdivided in serially homologous units, the so-called segments. In many arthropods, ventral and dorsal segmental tissue typically is aligned in parallel, but is dependent on different genetic inputs. In the pill millipede Glomeris marginata (Myriapoda: Diplopoda), ventral and dorsal segmental patterning is clearly decoupled providing an excellent model for the investigation of ventral versus dorsal segmentation mechanisms. This paper reports on the finding of a single embryo that lacks dorsal segmental and extraembryonic tissue. Ventral derivatives, however, are widely developed normally. This suggests that ventral and dorsal tissue is not only patterned differently, as shown previously, but also that ventral tissue can develop (or at least persist) independently from dorsal tissue. It also suggests a correlation of dorsal segmentation and function of the extraembryonic tissue. This assumed correlation may involve the guidance of the two dorsal hemispheres of the developing embryo dorsally, or that formation and/or maintenance of extraembryonic tissue depends on the input of dorsal segmental tissue. Whether the observed abnormalities are caused by mutation or are the result of otherwise disturbed early development is unclear.

The first records of mass occurrences (swarming behaviour) in giant pill-millipedes, order Sphaerotheriida, are reported from Madagascar. Swarming behaviour in the order Sphaerotheriida seems to be restricted to a single of more than sixty described Malagasy species, Zoosphaerium neptunus (Butler, 1872), which is the world’s largest known giant pill-millipede. Rolled-up individuals can be up to the size of a baseball, tennis ball or small orange, but only females reach this giant size, males being smaller than a ping-pong ball. Nine occurrences of such Z. neptunus swarms were analyzed based on actual specimens, video or photographic evidence collected by other researchers, dating back as far as 1892. One additional swarm, comprising several thousand individuals was examined in detail, with 260 randomly collected specimens being dissected and measured. The findings highlight that the swarming behaviour in Z. neptunus differs from that of all other millipedes in two important details: (1) The individual swarm is restricted to specimens of a single related size (and presumably age) class; often only sexually immature individuals; (2) the swarming behaviour is obligate, most, if not all specimens of the species in a given area participate in such swarms; Z. neptunus specimens are rarely, if ever, not found in a swarm. Reasons behind such massing events in millipedes are currently little understood, but a potential explanation for the mass occurrences in Z. neptunus might be higher survival rate from predation in combination with a close sibling relationship between members of one swarm. The almost obligate swarming behaviour in the widespread Z. neptunus species might represent a conservation problem, since whole swarms, and therefore a whole generation in a given area, can be lost through anthropogenic interferences such as over-collecting for the pet trade, habitat fragmentation or road kills. RESUME Il est rare que l’Homme se retrouve au contact des myriapodes et plus rare encore que de cette rencontre naisse un antagonisme. On peut citer le cas du mille-pattes indien Xenobolus carnifex qui degrade les toits et celui des essaimages de myriapodes qui entrainent souvent de graves problemes dont le plus courant est l’invasion de maisons, voire de villages entiers. Ce comportement inhabituel est surtout mentionne pour l’Europe et les raisons de ce gregarisme chez les mille-pattes sont encore peu comprises. Certains pensent que ces rassemblements sont lies a la surpopulation ou qu’ils augmentent l’efficacite des defenses chimiques. Une des rares similitudes entre ces phenomenes est que les specimens y participant sont adultes ou subadultes, jamais juveniles. Cet article fait etat de la premiere observation d’un regroupement en masse (comportement gregaire) chez les grands mille-pattes volveurs, ordre des Sphaerotheriida, a Madagascar. Au sein de cet ordre, le comportement gregaire semble se limiter a une seule espece parmi plus de soixante especes malgaches connues, Zoosphaerium neptunus (Butler, 1872), actuellement l’espece la plus grande des mille-pattes volveurs connus. Enroules, des specimens peuvent atteindre la taille d’une balle de baseball, de tennis, ou d’une petite orange ; mais cela ne concerne que les femelles, les mâles etant plus petits qu’une balle de ping-pong. Notre etude se fonde sur l’analyse de neuf cas d’un tel gregarisme chez Z. neptunus , soit a partir d’observations directes, soit a partir de documents, notamment photographiques ou video, recueillis par d’autres chercheurs, et ce aussi loin que 1892. Nous avons examine en detail un essaim compose de plusieurs milliers de specimens, dont 260 recoltes au hasard, disseques et mesures. Les resultats mettent en evidence que le comportement gregaire de Z. neptunus differe de celui des autres mille - pattes sur deux points importants : (1) chaque essaim ne comporte que des specimens de taille proche (et probablement d’un âge similaire), souvent exclusivement des specimens immatures sexuellement ; (2) le comportement gregaire est obligatoire ; la plupart, si ce n’est tous les specimens de l’espece d’une zone donnee participent a ce genre d’essaimages, - Z. neptunus est rarement, sinon jamais, observe hors d’un essaim. On ignore les causes du gregarisme chez les mille-pattes, mais une explication possible du regroupement en masse chez Z. neptunus pourrait etre un meilleur taux de survie face a la predation en combinaison avec une relation fraternelle renforcee entre les membres d’un meme essaim. Le comportement gregaire quasi-obligatoire chez le tres repandu Z. neptunus pourrait compliquer sa preservation du fait que les interferences humaines telles que sa surexploitation a des fins commerciales, la fragmentation de son habitat ou le risque de mourir sur la route peuvent entrainer la perte d’essaims entiers, et donc de toute une generation dans une zone donnee.

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.

What is a segment? In a thoughtful Perspective, [Minelli][1] explores segmentation during development of the pill millipede, and in so doing challenges prevailing views of what constitutes a segment. [1]: http://www.sciencemag.org/cgi/content/full/306/5702/1693

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.

AdultGlomeris marginata reloaded glandular defensive fluid slowly and fairly constantly at 15 °C for 100 days after being milked to depletion. Female millipedes produced more sticky exudate than males, but the two sexes stored secretion at approximately the same absolute rate, 12 μg/individual/day. Hence, males, which weighed one third as much as females, accumulated disproportionately more secretion. MaleGlomeris in the reloading treatments after 75 days yielded as much exudate as controls, millipedes given 100 days to supplement their field reserves, whereas experimental females always produced less secretion than controls. Projections from these data suggest that adultGlomeris, regardless of sex, require more than 4 months to replenish their defensive reserves after completely discharging them in an attack.

Numerous spiders mimic ants, and considerable descriptive literature is available (Bonnet, 1945); other spiders are supposed to resemble bird droppings or certain insects. On the other hand, literature on animals that mimic spiders is scant and no references could readily be found. Spiders, however, are poisonous to various groups of animals and should present evolutionary opportunities to potential mimics. Spider beetles (Ptinidae) at times are brought in, mistaken by collectors for spiders; their relationship to spiders is obscure. On a trip to Dalmatia, in the vicinity of Dubrovnik, I found the pill bug, Armadillidium klugii Brandt 1833 (Crustacea: Isopoda), to be so similar to Latrodectus mactans tredecimguttatus (Rossi, 1790), the malmignatte or European form of the black widow, as to be easily mistaken for it. The northern Mediterranean widows have bright red spots on a glistening black body (Fig. la), coloration resembling that of juvenile American forms. The spots are usually absent in adults in other parts of the range of L. mactans. The spiders are found among piles of stones, roadbanks, and deep plow furrows. Among stones in the Dubrovnik area we found Armadillidium klugii, a pill bug endemic in this area of Dalmatia and on adjacent islands, that rolls up when disturbed, showing bright red spots on a shiny black background (Fig. lc), strikingly like the abdomen of Latrodectus. The isopod is found commonly in crevices and between rocks. It was described from alcoholic specimens (Strouhal, 1928) in which the red color had washed out leaving yellow spots instead of red. Bright colors are not commonly found among pill bugs, and the coloration thus is most unusual. According to Strouhal, the species is variable in color, black to brown-black, with spots whitish to yellow and telson yellow to dark in alcohol. No black widows were collected with the isopod but this was perhaps due to the extremely low population of L. mactans in summer of 1962, the lowest in at least 10 years both in Italy and Yugoslavia (personal communication, Z. Maretic and M. Alessandrini), possibly due to unusually cold weather late in spring. The area probably normally supports a large black-widow population. There also occurs on the ground between stones in the Dubrovnik area, a pill millipede Glomeris pulchra C. L. Koch 1847 (Diplopoda: Glomerida), a black species with a series of bright red spots. When disturbed the millipede rolls up like a pill bug (Fig. lb), also strikingly resembling the malmignatte abdomen. Dr. P. Tongiorgi made me aware that pill millipedes of the genus Glomeris in northern Italy may have a similar color pattern. Live animals of a different species from Pisa, Italy, were also all black with two series of spots varying in color from yellowish to reddish; none had bright red spots, but no specimens were obtained from areas where Latrodectus is common.' Mimicry depends upon a characteristic of a model that makes it of selective advantage to be confused with the model. Latrodectus is known to be extremely poisonous to some vertebrates-guinea pigs, horses, camels, and man. Its poison must be effective on some invertebrates as under its webs were found scarabid beetles several times the volume of Latrodectus. The nocturnal habit of mammals and their sensitivity to the poisons would preclude any advantage of mimicry as a means of avoiding predation by them. Superficial acquaintance with the habitat suggests that lizards, abundant everywhere, might be the most important predators of isopods, glomerid millipedes, and ground-dwelling spiders. The only reference that could be found (Vellard, 1936) indicates that Latrodectus is poisonous, but not lethal, to lizards. If the spider should succeed in biting a lizard attempting predation, the lizard would have an opportunity to learn not to attack Latrodectus again, and mimicry might very well give a selective advantage against lizard predation. As far as is known (Mertens, personal communication), Mediterranean lizards of the genus Lacerta feed on isopods, and it is believed that the rolling up of Armadillidium would not serve as complete protection. Another possibility is that the mimics display similar aposematic coloration. However, isopods are not known to be distasteful and the defensive secretions of most millipedes are not well-developed in the glomerids; a pair of glands is present but the secretions are odorless and tasteless (Attems, 1926).