Two new species of the millipede family Blaniulidae (Diplopoda, Julida) from caves in central and southeastern Europe.

Two new species, one each of the previously monotypic genera Cibiniulus Verhoeff, 1927, and Thassoblaniulus Mauriès, 1985, are described from caves in Slovakia and Albania, respectively: C. slovacus Antic, Mock & Enghoff, sp. n. and T. radjai Antic & Enghoff sp. n. Brief discussions on relationships with congeners and on the taxonomic position of the genera Cibiniulus and Thassoblaniulus within Blaniulidae are presented, as well as a distribution map for both genera.

Molecular phylogeny of the Thyropygus allevatus group of giant millipedes and some closely related groups.

Giant cylindrical millipedes of the family Harpagophoridae, especially species of the genus Thyropygus, are broadly distributed in Thailand and nearby countries. They show a great deal of variation in body size, color patterns and gonopodal characters. Phylogenetic analyses of 26 nominate species from six genera in the subfamilies Harpagophorinae and Rhynchoproctinae, as well as nine new morphotypes (regarded as new species), were performed with the DNA sequences from two mitochondrial gene fragments (16S rRNA and COI). The genus Thyropygus (Harpagophorinae) was recovered as monophyletic under all analyses, whilst the representatives of Rhynchoproctinae also formed a monophyletic group. However, the analyses suggested that the tribe Gonoplectini should be elevated to a separate subfamily, Gonoplectinae. The molecular analyses were largely (but not totally) congruent with, and so supported the usefulness of, gonopodal characters for the classification and identification of harpagophorid millipedes, and additionally supported previous studies on the delimitation of species and subgroups. This is the first molecular study inside the family Harpagophoridae and provides the basis for further studies of the evolutionary processes and biogeographic patterns of millipedes in Southeast Asia.

Segmentation of the millipede trunk as suggested by a homeotic mutant with six extra pairs of gonopods.

BACKGROUND: The mismatch between dorsal and ventral trunk features along the millipede trunk was long a subject of controversy, largely resting on alternative interpretations of segmentation. Most models of arthropod segmentation presuppose a strict sequential antero-posterior specification of trunk segments, whereas alternative models involve the early delineation of a limited number of 'primary segments' followed by their sequential stereotypic subdivision into 2n definitive segments. The 'primary segments' should be intended as units identified by molecular markers, rather than as overt morphological entities. Two predictions were suggested to test the plausibility of multiple-duplication models of segmentation: first, a specific pattern of evolvability of segment number in those arthropod clades in which segment number is not fixed (e.g., epimorphic centipedes and millipedes); second, the occurrence of discrete multisegmental patterns due to early, initially contiguous positional markers. RESULTS: We describe a unique case of a homeotic millipede with 6 extra pairs of ectopic gonopods replacing walking legs on rings 8 (leg-pairs 10-11), 15 (leg-pairs 24-25) and 16 (leg-pairs 26-27); we discuss the segmental distribution of these appendages in the framework of alternative models of segmentation and present an interpretation of the origin of the distribution of the additional gonopods.The anterior set of contiguous gonopods (those normally occurring on ring 7 plus the first set of ectopic ones on ring 8) is reiterated by the posterior set (on rings 15-16) after exactly 16 leg positions along the AP body axis. This suggests that a body section including 16 leg pairs could be a module deriving from 4 cycles of regular binary splitting of an embryonic 'primary segment'. CONCLUSIONS: A very likely early determination of the sites of the future metamorphosis of walking legs into gonopods and a segmentation process according to the multiplicative model may provide a detailed explanation for the distribution of the extra gonopods in the homeotic specimen. The hypothesized steps of segmentation are similar in both a normal and the studied homeotic specimen. The difference between them would consist in the size of the embryonic trunk region endowed with a positional marker whose presence will later determine the replacement of walking legs by gonopods.

Laboulbeniales on millipedes: the genera Diplopodomyces and Troglomyces.

The genera Diplopodomyces and Troglomyces (Laboulbeniales, Ascomycota) parasitizing millipedes (Diplopoda) are reviewed. We describe four new species: D. lusitanipodos parasitic on Lusitanipus from Portugal and on Cyphocallipus from Spain, D. veneris on Lusitanipus from Portugal, T. botryandrus on Leptoiulus from Italy and T. rossii on Ophyiulus, Leptoiulus and Typhloiulus from Italy. Diplopodomyces veneris is a dimorphic species, with different morphotypes growing on the male copulatory organs and on the surroundings of the female gonopore respectively. The type species of both genera also have been studied, their descriptions are revised and a neotype for T. manfrediae is designated. New records from Italy are given for D. callipodos and T. manfrediae.

Insular species swarm goes underground: two new troglobiont Cylindroiulus millipedes from Madeira (Diplopoda: Julidae).

Two new species of the genus Cylindroiulus Verhoeff, 1894, C. julesvernei and C. oromii, are described from the subterranean ecosystem of Madeira Island, Portugal. Species are illustrated with photographs and diagrammatic drawings. The new species belong to the Cylindroiulus madeirae-group, an insular species swarm distributed in the archipelagos of Madeira and the Canary Islands. We discuss the differences between the new species and their relatives and present information on the subterranean environment of Madeira. An updated overview of the subterranean biodiversity of millipedes in Macaronesia is also provided.

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 cave-dwelling millipede of the genus Scutogona from central Portugal (Diplopoda, Chordeumatida, Chamaesomatidae).

A new cave-dwelling species of the genus Scutogona Ribuat, 1913, S. minor n. sp., is described from caves of Sicó karst in central Portugal. The classification and delimitation of Scutogona vis-à-vis related genera, in particular Meinerteuma Mauriès, 1982, is discussed.

The genus Boreviulisoma Brolemann, 1928--an Iberian-N African outlier of a mainly tropical tribe of millipedes (Diplopoda: Polydesmida: Paradoxosomatidae.

The genus Boreviulisoma Brolemann, 1928, is revised. The synonymy of Liliputia Attems, 1952, under Boreviulisoma is confirmed, but L. badia Attems, 1951, from Spain, is resurrected, as Boreviulisoma badium, from synonymy under B. liouvillei Brolemann, 1928 (the type species of Boreviulisoma) from Morocco. Boreviulisoma barrocalense n. sp. is described from the subterranean environment of the Algarve, the southernmost province of Portugal. The distribution of the three known species of Boreviulisoma is mapped, main characters of the genus and its species are reviewed and a key to species of the genus is included. The isolated occurrence of Boreviulisoma badium and B. barrocalense n. sp. in the southern Iberian Peninsula, together with B. liouvillei and Eviulisoma abadi Mauries, 1985, in Morocco, as northern outposts of the chiefly Afrotropical-Neotropical tribe Eviulisomatini, is discussed.

Subterranean species of Acipes Attems, 1937 (Diplopoda, Julida, Blaniulidae).

Two new blind, cave-dwelling species of the genus Acipes Attems, 1937, are described from the Algarve, southern Portugal: A. machadoi n. sp. and A. bifilum n. sp. Acipes andalusius Enghoff & Mauriès, 1999, is reported from the mesovoid shallow substratum in Alicante (Spain), 250 km from the type locality in Andalusia.

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.

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.

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.

Phylogenetic relationships in the millipede family Julidae.

A phylogenetic analysis of 40 species (22 genera) of the Palaearctic millipede family Julidae was made based on partial sequences of the mitochondrial 16S rRNA (16S) gene and the nuclear 28S rRNA (28S) gene, respectively. The two data sets (16S rDNA and 28S rDNA) were analysed individually and in combination using direct optimization as implemented in POY. The 16S rDNA and the 28S rDNA sequences vary from 410 to 449 bp and from 467 to 525 bp in length, respectively. All searches were performed under six different gap opening costs, an extension gap cost of 1, and a substitution cost of 2. Incongruence length difference values were used to select the preferred tree. The order Julida was recovered as monophyletic under all weight sets. The family Julidae was recovered as monophyletic except under one weight set where the genus Nepalmatoiulus is sister to all other Julida. Within Julidae, a clade of Paectophyllini + Calyptophyllini is sister to all others on the preferred tree but this relationship is not robust. A hitherto unrecognized clade of (South) east Asian genera (Anaulaciulus and Nepalmatoiulus) was recovered under five weight sets. Another "new" robust clade (Oncoiulini + Schizophyllini) is congruent with a hitherto unrecognized complex morphological character. Further clades recovered within the Julidae partly conflict with the accepted classification, which is only to a limited extent based on phylogenetic arguments.

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.

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.

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.

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.

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.