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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 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.

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.

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.

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.

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).