Phylogenetic relationships of millipedes in the subclass Penicillata (Diplopoda) with a key to the genera

Bristly millipedes (subclass Penicillata, order Polyxenida) are minute diplopods characterised by uncalcified cuticle and a body covered with unique tufts of bristles. The order is found worldwide and comprises less than 200 described species divided into three families, with many of the species and genera being poorly known. The first evolutionary analysis of the order presented here utilises both molecular (COI, 18S rRNA and 16S rRNA) and morphological data to examine monophyly of the families and subfamilies and the evolutionary relationships between them. Maximum likelihood analysis was based on molecular data only, whereas parsimony analyses were based on molecular data as well as combined morphological and molecular data. The results of these analyses with two different optimality criteria were incongruent in many aspects. Unlike parsimony, the likelihood result found strong support for a basal position of the family Synxenidae and separation of the order into two monophyletic clades corresponding to the two superfamilies Synxenoidea, containing the family Synxenidae, and Polyxenoidea, containing the families Polyxenidae and Lophoproctidae. Parsimony results did not support the existence of the two superfamilies. Both analyses resolved the family Synxenidae as monophyletic and Polyxenidae as polyphyletic, whereas the family Lophoproctidae was shown to be paraphyletic in likelihood and monophyletic in parsimony analysis. The subfamilies Monographinae and Polyxeninae were found to be monophyletic in the likelihood tree but parsimony suggested paraphyly of both. The results suggest that further revision of the systematics of the Polyxenida may be necessary. However, a much larger molecular data set will be necessary to clarify and provide stronger nodal support for phylogenetic trees and to confirm the relationships, particularly of the families Polyxenidae and Lophoproctidae. Molecular identification is likely to be an important tool for this taxonomically challenging order in future. A simplified key to the genera is provided.     

Revision of the millipede genus Parafontaria Verhoeff, 1936 (Diplopoda,Xystodesmidae)

The xystodesmid millipede genus Parafontaria Verhoeff, 1936 is revised in the light of the geographic variation found in the genital morphology of both sexes. The following 11 species, including one new species, are recognized as valid: P. erythrosoma (Takakuwa, 1942), P. ishiii Shinohara, 1986, P. doenitzi (Karsch, 1880), P. laminata (Attems, 1909), P. crenata Shinohara, 1986, P. longa Shinohara, 1986, P. tokaiensis n. sp., P. shiraiwaensis Shinohara, 1986, P. falcifera (Verhoeff, 1936), P. spathulata (Miyosi, 1951) and P. takakuwai (Shinohara, 1957). Parafontaria tonominea (Attems, 1899) can be recognized as a species complex and is operationally defined as a species. The systematic status of P. terminalis (Takakuwa, 1942) is uncertain. The following new synonymies are proposed: Fontaria (Parafontaria) armigera Verhoeff, 1936 (currently P. laminata armigera (Verhoeff, 1936)), F. (P.) kuhlgatzi Verhoeff, 1937 (currently P. kuhlgatzi (Verhoeff, 1937)) and P. echizenensis Shinohara, 1986 with P. laminata; F. coarctata circula Attems, 1901 (currently P. circula (Attems, 1901)), F. coarctata acutidens Attems, 1909 (currently P. acutidens (Attems, 1909)), F. (Japonaria) spiraligera Verhoeff, 1937 (currently P. spiraligera (Verhoeff, 1937)), F. (J.) marmorata Verhoeff, 1937 (currently P. marmorata (Verhoeff, 1937)), Japonaria longispinosa longispinosa Miyosi, 1951 (currently P. longispinosa longispinosa (Miyosi, 1951)), J. longispinosa falcata Miyosi, 1951 (currently P. longispinosa falcata (Miyosi, 1951)), J. egregia Haga, 1968 (currently P. egregia (Haga, 1968)) with P. tonominea. A cladistic analysis based on 15 morphological characters was performed for all species except P. terminalis. The strict and majority rule (50%) consensus trees identified three and five clades, respectively, within the genus.     

A new cave‐dwelling millipede of the genus Bollmania Silvestri, 1896 from Yunnan,China, with remarks on the reduction of the second female leg‐pair (Diplopoda: Callipodida: Caspiopetalidae)

Bollmania beroni sp. n., described from a cave in Jianshui County, Yunnan, China is the first true troglo‐ and hygrophilic species in the genus. The new locality extends the range of Bollmania ca 2500?km towards SE, ca 1700?km SW of the only other Chinese record. Notes are given on B. orientalis (Silvestri, ), B. nodifrons Lohmander, and B. oblonga Golovatch, , based on new material from Turkmenistan and Tajikistan, and on two unnamed, probably new species from Afghanistan. An updated key to the eight described species is presented. Original observations and illustrations of second female legs in various callipodid genera are presented, along with a literature review of this character, which has so far received little attention from taxonomists.     

Size criteria for the distinction between Cylindroiulus londinensis (Leach) and Cylindroiulus caeruleocinctus (Wood) (Diplopoda: Julidae)

Ring number, body height and live weight are given for each stadium and sex of two Cylindroiulus species often confused in the literature. In each stadium there is no overlap between the height and weight of the two species, which makes it possible to identify specimens using size criteria alone. According to data from different places in Europe, intraspecific variation is not substantial enough to fill the size gap between the two species.     

Taxonomic review of the Amazonian millipede genus Parastenonia Hoffman, 1977 and description of a new species from iron-ore caves (Polydesmida: Chelodesmidae)

ABSTRACT

The Amazonian genus Parastenonia Hoffman, 1977 is here revised. The species P. carajas sp. nov. is described from Brazilian iron-ore caves and P. aurae (Schubart, 1947) is considered junior synonym of P. parae (Cook, 1895). In addition, the known geographic distribution of the genus is presented with the inclusion of new occurrence data.

http://www.zoobank.org/urn:lsid:zoobank.org:act:CCBEED10-DCB4-48C9-A71D-CED3462DD72A     

Monophyly of the ring-forming group in Diplopoda （Myriapoda, Arthropoda） based on SSU and LSU ribosomal RNA sequences

Two controversies exist in the phylogeny of the derived millipedes (Diplopoda). The first is whether millipedes with a fusion ring, including Polydesmida, Spirobolida, Spirostreptida and Julida, form a monophyletic group (the ring-forming group). The second concerns the phylogenetic relationship within the three orders of Juliformia, i.e. Julida, Spirostreptida and Spirobolida. To resolve these phylogenetic controversies, we sequenced 18S and 28S rDNA from six millipede orders and retrieved several homologous sequences from GenBank. Our results give robust support to the monophyly of the ring-forming group based on maximum parsimony methods, maximum likelihood methods and Bayesian inference. The monophyly of the ring-forming group suggests that the fusion of segment sclerites might have occurred only once during millipede evolutionary history. We also established a sister-group relationship between Spirobolida and Spirostreptida within Juliformia after eliminating a short-branch attraction phenomenon, which is consistent with that from the mitochondrial genome analysis.     

Revision of the millipede genus Xystodesmus,with reference to the status of the tribe Xystodesmini (Diplopoda: Xystodesmidae)

On the basis of morphometrics and geographic distribution, we revise Xystodesmus martensii Peters and its related forms in Japan, which had been recognized as ten species belonging to five genera. In delimitation of species, first we sought the biggest pheno-geographic units based on the dendrogram from a cluster analysis and geographic distribution; a pheno-geographic unit is defined as an aggregation of populations being phenetically closely similar and geographically closely distributed to each other. Then, we decided whether each unit was warranted as a species. As a result, we propose that these forms constitute a single genus Xystodesmus comprising six species including one new species. The newly defined Xystodesmus is characterized by high variation in gonopod morphology and conservative coloration, body form and size. The status of some populations is still uncertain because of lack of material. The tribe Xystodesmini, to which Xystodesmus belongs, and the tribe Harpaphini are so similar that we could not warrant their separation. We propose the following new synonymies: the tribe Harpaphini with the tribe Xystodesmini; Phrurodesmus Takakuwa and Nikkonus Chamberlin and Wang with Xystodesmus; Xystodesmus scabra Verhoeff, Rhysodesmus tuberculatus Takakuwa, Rh. ikaoensis Takakuwa, and Rh. kitazawai Miyosi, with X. martensii; Phrurodesmus kinshaensis Murakami with X. gracilipes (Takakuwa). Riukiaria shirozui Takakuwa, Rh. serrulatus Miyosi, and P. gracilipes Takakuwa, are transferred to Xystodesmus. We describe X. tokaiensis as a new species.     

A second species of the genus Vietnamorpha Golovatch, 1984 (Polydesmida: Paradoxosomatidae) and notes on the generic relationship

A second Vietnamorpha species is described from Pu Mat National Park, north-central Vietnam. The new species is distinguished by the gonopod femorite being erect and cylindrically elongate; and the solenomere being completely sheathed by the solenophore, and being only exposed at the tip. The relationship between Vietnamorpha and several sulciferinine genera was analysed using a combination of a mitochondrial gene (16S rRNA) and two nuclear genes (18S and 28S rRNA). The genus Vietnamorpha is more closely related to the genus Anoplodesmus than to other sulciferinine genera.

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