A new species of Lophostreptus Cook, 1895 discovered among syntypes of L.regularis Attems, 1909 (Diplopoda, Spirostreptida, Spirostreptidae).

A new species of the genus Lophostreptus Cook, 1895 is described, based on specimens hidden for over a century among the syntypes of its congener Lophostreptusregularis Attems, 1909 housed in the Naturhistoriska Riksmuseet Stockholm (NRMS) and the Naturhistorisches Museum Wien (NHMW). A lectotype is designated for Lophostreptusregularis Attems, 1909 in order to stabilize its taxonomy. Updates to the millipede fauna of Mt. Kilimanjaro, Tanzania are provided.

A new distinct, disjunct giant millipede of the genus Spirostreptus Brandt, 1833, from Tanzania, and a solution for orphaned Spirostreptus species (Diplopoda, Spirostreptida, Spirostreptidae).

Spirostreptus digitus sp. nov. is described based on specimens from Tanzania, and the genus Spirostreptus Brandt, 1833 is discussed, as is the higher classification of the family Spirostreptidae. Gonopod illustrations of S. heros Porat, 1872, and S. tripartitus Cook & Collins, 1893, are provided for comparison with the new, morphologically very distinct species. Analocostreptus Silvestri, 1910, is suggested as the valid genus name for the species currently being placed in Spirostreptus sensu auctorum, nec Brandt, 1833, and 32 nominal species of Spirostreptus are placed in new combinations with Analocostreptus, viz., Analocostreptus amandus (Attems, 1914), comb. nov., Analocostreptus biconus (Attems, 1934), comb. nov., Analocostreptus bonifatius (Attems, 1914), comb. nov., Analocostreptus castaneus (Attems, 1934), comb. nov., Analocostreptus cornutus (Attems, 1934), comb. nov., Analocostreptus damasus (Attems, 1953), comb. nov., Analocostreptus dartevellei (Attems, 1953), comb. nov., Analocostreptus dentiger (Attems, 1953), comb. nov., Analocostreptus garambanus (Chamberlin, 1927), comb. nov., Analocostreptus gregorius (Attems, 1914), comb. nov., Analocostreptus ineptus (Kraus, 1958), comb. nov., Analocostreptus informis (Attems, 1938), comb. nov., Analocostreptus makarius (Attems, 1914), comb. nov., Analocostreptus manyemanus (Attems, 1927), comb. nov., Analocostreptus manyemanus biserialis (Attems, 1938), comb. nov., Analocostreptus medjensis (Chamberlin, 1927), comb. nov., Analocostreptus micromelas (Saussure & Zehntner, 1902), comb. nov., Analocostreptus missionarius (Attems, 1953), comb. nov., Analocostreptus montivagus (Karsch, 1881), comb. nov., Analocostreptus multisulcatus (Demange, 1957), comb. nov., Analocostreptus pancratius (Attems, 1914), comb. nov., Analocostreptus pavani (Demange, 1981), comb. nov., Analocostreptus phthisicus (Saussure & Zehntner, 1902), comb. nov., Analocostreptus pictus (Saussure & Zehntner, 1902), comb. nov., Analocostreptus rolini (Silvestri, 1897), comb. nov., Analocostreptus sculptus (Saussure & Zehntner, 1902), comb. nov., Analocostreptus semilunaris (Peters, 1855), comb. nov., Analocostreptus servatius (Attems, 1914), comb. nov., Analocostreptus tetricus (Attems, 1934), comb. nov., Analocostreptus tiburtius (Attems, 1953), comb. nov., Analocostreptus triangulicollis (Attems, 1934), comb. nov., Analocostreptus versicolor (Saussure & Zehntner, 1902), comb. nov. and Analocostreptus yambatanus (Attems, 1934), comb. nov. Illustrations of the gonopods of A. ibanda (Silvestri, 1907), type-species of Analocostreptus, are provided, and the species is recorded as new for Kenya.

Redescription of the giant Southeast Asian millipede Spirobolusmacrurus Pocock, 1893 and its assignment to the new genus Macrurobolus gen. nov. (Diplopoda, Spirobolida, Pachybolidae).

A new genus of the millipede family Pachybolidae from Southeast Asia is described: Macrurobolus gen. nov., with Spirobolusmacrurus Pocock, 1893 as type species. This latter species is DNA barcoded (COI) and redescribed based on male morphological characters, which hitherto were unknown. The new genus differs from other pachybolid genera by having (1) the preanal ring process long and protruding beyond the anal valves and (2) the anterior gonopod telopodite distally abruptly narrowed, forming an extremely long, slender, elevated process curved caudad. Given that Macrurobolus gen. nov. is a monotypic genus, it is aphyletic and thus requires further taxonomic revision.

A new species of the hitherto monospecific genus Pleonoporus Attems, 1938 (Diplopoda, Spirostreptida, Odontopygidae).

The hitherto monospecific genus Pleonoporus is revised based on the syntypes of P.robustus Attems, 1938, housed in Naturhistorisches Museum Wien (NHMW), as well as on specimens of what we interpret as a new species of the genus, Pleonoporustanzanicus sp. nov., collected in Tanzania and housed in the Museum of Nature - Zoology, Leibnitz Institute for the Analysis of Biodiversity Change (ZMH) for more than a century. Both species are described based on habitus and gonopod structures and illustrated with micrographs, whereas scanning electron microscope images are also provided for the new species. This paper further highlights the importance of natural history collections for taxonomic research and studies on species diversity in general.

Two new millipede species of the genus Coxobolellus Pimvichai, Enghoff, Panha & Backeljau, 2020 (Diplopoda, Spirobolida, Pseudospirobolellidae).

Two new millipede species of the genus Coxobolellus Pimvichai, Enghoff, Panha & Backeljau, 2020 from Thailand are described: Coxobolellussaratani sp. nov. from Loei Province and Coxobolellusserratoligulatus sp. nov. from Uttaradit Province. The descriptions are based on gonopod morphology and two mitochondrial gene fragments (COI and 16S rRNA). The phylogenetic mtDNA analysis assigned the two new species unequivocally to the consistently well-supported Coxobolellus clade, in which they form a fifth subclade that was well supported by maximum likelihood analysis of 16S rRNA, though neither by Bayesian inference nor by COI. The two new Coxobolellus species share four conspicuous gonopodal synapomorphies of the genus: (1) the protruding process on the coxae of the 3rd (and sometimes 4th) pair of male legs, (2) a large, triangular coxae on the 4th-5th pair of legs, (3) a short process of the preanal ring protruding as far as, or slightly beyond, the anal valves, and (4) the posterior gonopod telopodite divided into two parts, with a conspicuous pore opening at the mesal margin at the end of the coxal part of the posterior gonopod. Thus, the two new species provide further evidence of the well-defined monophyly of the genus Coxobolellus. Finally, the paper provides an updated morphological identification key to all currently described Coxobolellus species.

Comparative morphological study of the gnathochilarium of millipedes of the suborder Cambalidea (Juliformia: Spirostreptida) assessed by discrete and morphometric character approaches

Phylogenetic relationships of the suborder Cambalidea have been a well-known focus of discussion over the past 100 years. The most recent approach to the suborder is based on the morphology of the sexual structure of males and of the gnathochilarium. The phylogenetic interpretation of the gnathochilarium has been especially criticized due to the poorly understood evolution of its components since no hypothesis of homology has been put forward. This work provides a comparative study of characters of the gnathochilarium. Twenty-one characters are proposed and optimized for 34 terminals from the orders Julida and Spirostreptida. Two datasets were analysed on the basis of whether the sclerite duplomentum corresponds to a mentum fused to a promentum or only to an elongated mentum (promentum being absent). Concatenated analyses were performed with both discrete and morphometric characters to verify the variation in shape of the mentum. In all topologies obtained, Cambalidea were recovered as paraphyletic in terms of the suborder Spirostreptidea. Through constrained searches, Cambalidae, Cambalopsidae and Iulomorphidae were not recovered, while Pseudonannolenidae were recovered only when Physiostreptinae were excluded. The morphological evidence indicates the convergence of the gnathochilarium in Spirostreptidea and in some members of Cambalidea, with independent reversions of the promentum.

Penetrative and non-penetrative interaction between Laboulbeniales fungi and their arthropod hosts.

Laboulbeniales are a highly specialized group of fungi living only on arthropods. They have high host specificity and spend their entire life-cycle on an arthropod host. Taxonomic characters of Laboulbeniales are based on the architecture of the cells of the parenchymal thallus, i.e. the visible part of the fungus outside the host. The extent of the fungus spreading inside the host-the haustorium-remains largely unknown. The attachment to the arthropod host is fundamental to understand the fungus-animal interaction, but how this truly occurs is unclear. Recent evidences question the strictly parasitic life-style of Laboulbeniales. We used micro-computed tomography (µCT) and 3D reconstructions to visualize, for the first time, the complete structure of Laboulbeniales species in situ on their hosts. We compared the haustoriate species, Arthrorhynchus nycteribiae on an insect host to the non-haustoriate species, Rickia gigas on a millipede host. Our results confirm that some Laboulbeniales species are ectoparasitic and have a haustorial structure that penetrates the host's cuticle, while others are ectobionts and are only firmly attached to the host's cuticle without penetrating it. The presence and the morphology of the haustorium are important traits for Laboulbeniales evolution, and key factors for future understanding of host dependence and specificity.

A new giant species of the millipede genus emPrionopetalum/em Attems, 1909 from Mt. Kilimanjaro, Tanzania (Diplopoda, Spirostreptida, Odontopygidae).

A new large species of the millipede genus Prionopetalum Attems, 1909, P. nessiae sp. nov. from Mt. Kilimanjaro, Tanzania, is described, with SEM imaging and illustrations of relevant taxonomic characters, and mapped. Comments on the significance of sternum 9 in descriptions of Odontopygidae, and on the intrageneric differences in Prionopetalum, are included.

The first Laboulbeniales (Ascomycota, Laboulbeniomycetes) from an American millipede, discovered through social media.

Laboulbeniales are highly specialized arthropod-associated fungi. The majority of the almost 2200 known species live on insects, although they also occur on other arthropod hosts. Recently, the number of Laboulbeniales associated with millipedes has increased considerably. Here we describe the first species of a Laboulbeniales fungus, Troglomyces twitteri sp. nov., from an American millipede. The new species was initially discovered on a photo of Cambala annulata (Say, 1821) from Ohio, USA, which had been shared on Twitter. A subsequent microscopic study of Cambala millipedes in museum collections in Denmark and France confirmed the discovery.

Description of the female morphology of the relict troglobiont millipede Cantabrodesmus lorioli Mauriès 1971 (Diplopoda, Polydesmida, Chelodesmidae), and new records in the deepest pit of Spain.

The troglobiont millipede, Cantabrodesmus lorioli Mauriès, 1971, inhabits a few caves in a small region of Northern Spain. It is considered a relict species because of its geographical isolation compared to others in its family, and its to some extent uncertain systematic position. Cantabrodesmus lorioli has been placed in three different families, and now is considered a member of the Chelodesmidae, a family with only one other known native European representative. There are few records of this species, and almost nothing is known about its biology. The only available information on this species is the description of the male and a list of caves where it has been captured. In the present work, we provide the first description of the female morphology, illustrated by scanning electron microscopy, and a series of new records from caves in a region where it had never been captured before, including the deepest known pit in Spain, Torca del Porrón. The female morphology does not show many differences from that of the male, but several characters are discussed. All of the known information on the morphology of the species is used to discuss the implications for its biology and systematic position.

Phylogenetic placement of the monotypic genus Holopodostreptus Carl, 1913 and notes on the systematics of Pseudonannolenidae (Spirostreptida : Cambalidea)

The genus Holopodostreptus Carl, 1913 is a poorly known monotypic genus endemic to Ecuador. It differs from most other Pseudonannolenidae by a set of characteristics of the gnathochilarium and gonopods. Herein, we provide descriptive notes of males of H. braueri Carl, 1913 based on recently collected material. Females of the species are described for the first time, and images of the vulvae are provided. In addition, we attempt for the first time to place Holopodostreptus in a phylogenetic context. With a total of 14 taxa and 41 morphological characters, and under implied weighting (k = 6), a single most-parsimonious tree is recovered (total fit = 35.86). The genus is recovered as sister-group of Physiostreptus Silvestri, 1903. The clade Holopodostreptus + Physiostreptus is supported by one synapomorphy (absence of setae on the prefemoral process of leg-pair 1 in males) and one homoplastic transformation (narrow telopodite). Based on the phylogenetic results, notes on the subfamilies Pseudonannoleninae, Cambalomminae and Physiostreptinae and a key for all pseudonannolenid genera are provided.

A revision of dragon millipedes IV: the new genus Spinaxytes, with the description of nine new species (Diplopoda, Polydesmida, Paradoxosomatidae).

Nine new species constituting the 'spiny' group of dragon millipedes are assigned to the new genus Spinaxytes Srisonchai, Enghoff & Panha, gen. n. Seven new species are described from Thailand: S.biloba Srisonchai, Enghoff & Panha, sp. n. and S.palmata Srisonchai, Enghoff & Panha, sp. n. from Surat Thani Province, S.hasta Srisonchai, Enghoff & Panha, sp. n. from Chumphon Province, S.krabiensis Srisonchai, Enghoff & Panha, sp. n. (type species) and S.sutchariti Srisonchai, Enghoff & Panha, sp. n. from Krabi Province, S.uncus Srisonchai, Enghoff & Panha, sp. n., and S.macaca Srisonchai, Enghoff & Panha, sp. n. from Phang Nga Province; as well as one from Malaysia, S.tortioverpa Srisonchai, Enghoff & Panha, sp. n., and one from Myanmar, S.efefi Srisonchai, Enghoff & Panha, sp. n. The new genus is endemic to South Myanmar, South Thailand, and Malaysia, and all new species are restricted to limestone habitats. All were exclusively found living on humid rock walls and/or inside small caves. Complete illustrations of external morphological characters, an identification key, and a distribution map are provided.

First continental troglobiont Cylindroiulus millipede (Diplopoda, Julida, Julidae).

The new species of millipede Cylindroiulusvillumi is described from a cave in the Estremenho karst massif in central Portugal. It is the first cave-adapted species of its genus with a strict subterranean life-style in continental Europe, and is the fifth blind species of the genus. The new species is illustrated with photographs and diagrammatic drawings. It is tentatively placed in the purely Iberian Cylindroiulusperforatus-group. The differences between the new species and its relatives are discussed, as well as its adaptations to a subterranean life-style.

Novelty upon novelty visualized by rotational scanning electron micrographs (rSEM): Laboulbeniales on the millipede order Chordeumatida.

Laboulbeniales are highly specific ectoparasitic fungi of arthropods (insects, millipedes, and arachnids). The first Laboulbeniales parasitizing the millipede order Chordeumatida (Diplopoda) were discovered and described as a new dioecious genus of Laboulbeniales, Thaxterimyces, to accommodate the new species T. baliensis. Also the millipede host is a new species and is described as Metopidiothrix sheari. This is the first time Laboulbeniales fungus and its millipede host are described as new together. Males of Metopidiothrix have the most extensive secondary sexual modifications in the entire class Diplopoda. Although nothing is known about the function of these modifications, the unique pattern of Laboulbeniales infection in the new millipede species is obviously related to host sexual behavior. Rotational Scanning Electron Micrographs (rSEM) are used to create a 3D comprehensive model to examine the fungal-host interaction, a more advanced visualization of the ectoparasitic fungus on its host. Laboulbeniales diversity on millipedes is still understudied, and a consistent effort is needed to unveil and understand the extent and diversity of this biological interaction. Due to their minute size and apparently non-detrimental effect on their hosts, Laboulbeniales in general have been largely ignored by mycologists and neglected by generations of entomologists. As a result a significant component of global biodiversity has been strongly underestimated, and a wealth of new discoveries is still to be made both in the field and in existing museum collections.

Pseudotibiozus Demange, 1970-millipedes of the Eastern Arc Mountains, Tanzania (Diplopoda, Spirostreptida, Spirostreptidae).

The genus Pseudotibiozus Demange, 1970, is discussed, its type species, P. cerasopus (Attems, 1914) is re-described based on type and new material, and P. zophoribates sp. nov. is described from the West Usambara Mountains, Tanzania. The millipede faunas of the West and East Usambara Mts are compared.

Out of the dark void: Ommatoiulus longicornis n. sp., a new julid from Spain (Diplopoda, Julida) with notes on some troglobiomorphic traits in millipedes.

A new millipede species, Ommatoiulus longicornis n. sp. is described from the subterranean habitat in eastern Spain, and the elongation of its antennae is discussed in relation to the habitat where it dwells. We take the occasion to discuss the elongation of appendages as a troglobiomorphic character in millipedes and shed the light on the various, sometimes controversial, hypotheses that could have led to these modifications in millipedes and other subterranean arthropods.

Catalogue of the type material in the entomological collection of the Natural History Museum of Denmark: basal hexapods.

A catalogue of type specimens of basal hexapods (Collembola, Protura, Diplura, Archaeognatha, Zygentoma) in the collection of the Natural History Museum of Denmark, University of Copenhagen is presented. The NHMD collection harbours type material of 175 species, and it is composed of 132 holotypes, 4 neotypes, 3 syntypes, 32 lectotypes, and 39 paratypes. For Diplura one neotype is newly designated. The order Protura is represented by 5 families, Collembola by 7 families, Diplura by 6 families, Archaeognatha by 1 family and Zygentoma by 2 families.

A revision of dragon millipedes I: genus Desmoxytes Chamberlin, 1923, with the description of eight new species (Diplopoda, Polydesmida, Paradoxosomatidae).

The dragon millipede genus Desmoxytes s.l. is split into five genera, based on morphological characters and preliminary molecular phylogenetic analyses. The present article includes a review of Desmoxytes s.s., while future articles will deal with Hylomus Cook and Loomis, 1924 and three new genera which preliminarily are referred to as the 'acantherpestes', 'gigas', and 'spiny' groups. Diagnostic morphological characters of each group are discussed. Hylomus is resurrected as a valid genus and the following 33 species are assigned to it: H. asper (Attems, 1937), comb. n., H. cattienensis (Nguyen, Golovatch & Anichkin, 2005), comb. n., H. cervarius (Attems, 1953), comb. n., H. cornutus (Zhang & Li, 1982), comb. n., H. draco Cook & Loomis, 1924, stat. rev., H. enghoffi (Nguyen, Golovatch & Anichkin, 2005), comb. n., H. eupterygotus (Golovatch, Li, Liu & Geoffroy, 2012), comb. n., H. getuhensis (Liu, Golovatch & Tian, 2014), comb. n., H. grandis (Golovatch, VandenSpiegel & Semenyuk, 2016), comb. n., H. hostilis (Golovatch & Enghoff, 1994), comb. n., H. jeekeli (Golovatch & Enghoff, 1994), comb. n., H. lingulatus (Liu, Golovatch & Tian, 2014), comb. n., H. laticollis (Liu, Golovatch & Tian, 2016), comb. n., H. longispinus (Loksa, 1960), comb. n., H. lui (Golovatch, Li, Liu & Geoffroy, 2012), comb. n., H. minutuberculus (Zhang, 1986), comb. n., H. nodulosus (Liu, Golovatch & Tian, 2014), comb. n., H. parvulus (Liu, Golovatch & Tian, 2014), comb. n., H. phasmoides (Liu, Golovatch & Tian, 2016), comb. n., H. pilosus (Attems, 1937), comb. n., H. proximus (Nguyen, Golovatch & Anichkin, 2005), comb. n., H. rhinoceros (Likhitrakarn, Golovatch & Panha, 2015), comb. n., H. rhinoparvus (Likhitrakarn, Golovatch & Panha, 2015), comb. n., H. scolopendroides (Golovatch, Geoffroy & Mauriès, 2010), comb. n., H. scutigeroides (Golovatch, Geoffroy & Mauriès, 2010), comb. n., H. similis (Liu, Golovatch & Tian, 2016), comb. n., H. simplex (Golovatch, VandenSpiegel & Semenyuk, 2016), comb. n., H. simplipodus (Liu, Golovatch & Tian, 2016), comb. n., H. specialis (Nguyen, Golovatch & Anichkin, 2005), comb. n., H. spectabilis (Attems, 1937), comb. n., H. spinitergus (Liu, Golovatch & Tian, 2016), comb. n., H. spinissimus (Golovatch, Li, Liu & Geoffroy, 2012), comb. n. and H. variabilis (Liu, Golovatch & Tian, 2016), comb. n.Desmoxytes s.s. includes the following species: D. breviverpa Srisonchai, Enghoff & Panha, 2016; D. cervina (Pocock,1895); D. delfae (Jeekel, 1964); D. des Srisonchai, Enghoff & Panha, 2016; D. pinnasquali Srisonchai, Enghoff & Panha, 2016; D. planata (Pocock, 1895); D. purpurosea Enghoff, Sutcharit & Panha, 2007; D. takensis Srisonchai, Enghoff & Panha, 2016; D. taurina (Pocock, 1895); D. terae (Jeekel, 1964), all of which are re-described based mainly on type material. Two new synonyms are proposed: Desmoxytes pterygota Golovatch & Enghoff, 1994, syn. n. (= Desmoxytes cervina (Pocock, 1895)), Desmoxytes rubra Golovatch & Enghoff, 1994, syn. n. (= Desmoxytes delfae (Jeekel, 1964)). Six new species are described from Thailand: D. aurata Srisonchai, Enghoff & Panha, sp. n., D. corythosaurus Srisonchai, Enghoff & Panha, sp. n., D. euros Srisonchai, Enghoff & Panha, sp. n., D. flabella Srisonchai, Enghoff & Panha, sp. n., D. golovatchi Srisonchai, Enghoff & Panha, sp. n., D. octoconigera Srisonchai, Enghoff & Panha, sp. n., as well as one from Malaysia: D. perakensis Srisonchai, Enghoff & Panha, sp. n., and one from Myanmar: D. waepyanensis Srisonchai, Enghoff & Panha, sp. n. The species can mostly be easily distinguished by gonopod structure in combination with other external characters; some cases of particularly similar congeners are discussed. All species of Desmoxytes s.s. seem to be endemic to continental Southeast Asia (except the 'tramp' species D. planata). Some biological observations (relationship with mites, moulting) are recorded for the first time. Complete illustrations of external morphological characters, an identification key, and distribution maps of all species are provided.

"Open access" growth histories in millipedes (Diplopoda).

A unique pattern of missing defence glands on certain body rings is described for two species of the millipede family Mongoliulidae, order Julida: Ussuriiulus pilifer Golovatch, 1980, and Koiulus interruptus Enghoff et al., 2017. Based on the patterns of missing glands observed in recently collected samples of the two species, numbers of podous and apodous body rings in successive stadia of the postembryonic development can be inferred for each individual millipede, which in turn allows the reconstruction of pathways of anamorphosis in these species. The inferred numbers of body rings in developmental stadia are compared with actual numbers observed on additional samples, including the type series, of U. pilifer. The pattern of missing glands in the two mongoliulid species is compared with the pattern of missing glands typical of the entire millipede order Polydesmida.

A giant African millipede re-discovered, re-located and re-classified after 120 years (Diplopoda, Spirobolida, Pachybolidae).

The giant millipede Pachybolus tectus Cook, 1897, described from Zanzibar, East Africa, and never re-collected till now, is re-described based on newly collected specimens from Cameroon, West Africa. The species is transferred to the genus Pelmatojulus Saussure, 1860, and compared with the most similar congener, P. togoensis (Cook, 1897). The type specimen of P. tectus, which no longer exists, probably was mis-labelled. An overview of the distribution of Pelmatojulus species and records of the genus from Cameroon are given, including P. brachysternus (Cook, 1897) as new to the fauna of Cameroon. Pelmatojulus insignis (Saussure, 1859) and P. togoensis are recorded as new to the fauna of Ghana.