Phylogenomic inference of species and subspecies diversity in the Palearctic salamander genus Salamandra.

The salamander genus Salamandra is widespread across Europe, North Africa, and the Near East and is renowned for its conspicuous and polymorphic colouration and diversity of reproductive modes. The phylogenetic relationships within the genus, and especially in the highly polymorphic species S. salamandra, have been very challenging to elucidate, leaving its real evolutionary history and classification at species and subspecies levels a topic of debate and contention. However, the distribution of diversity and species delimitation within the genus are critically important for identifying evolutionarily significant units for conservation and management, especially in light of threats posed by the pathogenic chytrid fungus Batrachochytrium salamandrivorans that is causing massive declines of S. salamandra populations in central Europe. Here, we conducted a phylogenomic analysis from across the taxonomic and geographic breadth of the genus Salamandra in its entire range. Bayesian, maximum likelihood and network-based phylogenetic analyses of up to 4905 ddRADseq-loci (294,300 nucleotides of sequence) supported the distinctiveness of all currently recognised species (Salamandra algira, S. atra, S. corsica, S. infraimmaculata, S. lanzai, and S. salamandra), and all five species for which we have multiple exemplars were confirmed as monophyletic. Within S. salamandra, two main clades can be distinguished: one clade with the Apenninic subspecies S. s. gigliolii nested within the Iberian S. s. bernardezi/fastuosa; and a second clade comprising all other Iberian, Central and East European subspecies. Our analyses revealed that some of the currently recognized subspecies of S. salamandra are paraphyletic and may require taxonomic revision, with the Central- and Eastern-European subspecies all being poorly differentiated at the analysed genomic markers. Salamandra s. longirostris - sometimes considered a separate species - was nested within S. salamandra, consistent with its subspecies status. The relationships identified within and between Salamandra species provide valuable context for future systematic and biogeographic studies, and help elucidate critical evolutionary units for conservation and taxonomy.

Sex chromosome conservation, DMRT1 phylogeny and gonad morphology in diploid Palearctic green toads (Bufo viridis subgroup).

Due to the prevailing sex chromosome homomorphy and large genome size, the knowledge on sex determination systems, sex chromosomes and sex-determining genes in amphibians remains scarce. Using 3 cross-amplifying sex-linked microsatellite markers, we uncover sex determination systems and sex chromosomes in purebred, diploid Palearctic green toads (Bufo viridis subgroup), which had so far only been characterized in laboratory-bred hybrids. Our data support an XY system in B. balearicus, B. viridis and B. variabilis. While females show recombination, it is strongly suppressed (or not detectable) in males. Markers corroborate the largest chromosome pair 1 (homologous to linkage group 1 of Xenopus tropicalis) to represent the sex chromosomes in diploid species of the B. viridis subgroup (B. siculus, B. shaartusiensis, B. balearicus, B. turanensis, B. variabilis, B. viridis, and probably B. boulengeri). This chromosome harbors DMRT1, a key gene of the sexual pathway in deeply divergent animal taxa. However, our phylogenetic analysis of a 600-bp fragment of that gene in diploid green toad taxa reveals that X and Y alleles cluster by species and not by gametolog. This suggests that XY-sequence similarity stems from occasional XY recombination within DMRT1, and we preliminarily reject its role as the master sex determination gene, pending future extension of this evidence to the entire DMRT1 gene. We further create a chain of evidence, which supports the hypothesis that linkage group 1 of X. tropicalis appears to be maintained as the largest chromosome (1), and thus is homologous in anuran karyotype evolution from pipid to hylid, bufonid and ranid anurans.

Phylogeny of Geophilomorpha (Chilopoda) inferred from new morphological and molecular evidence.

To address the phylogenetic relationships of the centipede order Geophilomorpha (more than 1000 species), we have reinterpreted and expanded the knowledge on their morphological disparity, and have doubled the amount of molecular data available. We performed maximum parsimony and maximum likelihood analyses, using 195 phylogenetically informative morphological characters for 80 species, and DNA sequences of 28S, 18S, 16S rRNA and COI for up to 48 species. We found strong support for the monophyly of Geophilomorpha, the basal dichotomy between Adesmata and Placodesmata = Mecistocephalidae, and the basal dichotomy within Adesmata between two clades that are recognized here as superfamilies Himantarioidea and Geophiloidea. With respect to the families currently in use, Himantarioidea comprises three well supported clades corresponding to (i) Oryidae, (ii) Himantariidae, and (iii) Schendylidae s.l. including Ballophilidae; Geophiloidea comprises another three supported clades corresponding to (iv) a new family Zelanophilidae, (v) Gonibregmatidae s.l. including Eriphantidae and Neogeophilidae, and (vi) Geophilidae s.l. including Aphilodontidae, Dignathodontidae, Linotaeniidae, and Macronicophilidae.

Chilopoda Geophilomorpha of Europe: a revised list of species, with taxonomic and nomenclatorial notes.

An annotated list is provided for the genera and species of Chilopoda Geophilomorpha recorded from Europe, including Macaronesia. The list derives from a critical evaluation of all published information. All synonyms are also listed and all taxonomic and nomenclatorial novelties are discussed. Additionally, all available genus-group and species-group names are listed, together with type species and type localities respectively.To date, 452 available species-group names and 95 available genus-group names have been applied to European geophilomorphs, together with another 10 unavailable names. A total of 179 species in 37 genera are provisionally recognized here, but the actual taxonomic identity of 84 of these species is uncertain because their morphology is in completely or imprecisely known. Another 5 species have been recorded from European localities but probably are not established in the wild, and another 8 species have been reported probably only erroneously. We introduce the following 116 new synonymies: Algerophilus hispanicus (Meinert, 1870) [= Geophilus arago nicus Daday, 1889], Bothriogaster signata (Kessler, 1874) [= Notiphilus taeniatus C.L. Koch, 1847, = N. sanguineus C.L. Koch, 1847, = B. affinis Sseliwanoff, 1879, = B. meinerti Sseliwanoff, 1879], Clinopodes C.L. Koch, 1847 [= Poabius C.L. Koch, 1847], Clinopodes carinthiacus (Latzel, 1880) [= Geophilus flavidus styriacus Attems, 1895, = G. trebevicensis poschiavensis Verhoeff, 1934], C. flavidus C.L. Koch, 1847 [= Geophilus flavidus pachypus Verhoeff, 1942, = G. flavidus faitanus Verhoeff, 1943, = G. flavidus improvisus Verhoeff, 1943, = G. flavidus karamani Verhoeff, 1943, = G. flavidus sorattinus Verhoeff, 1951], Dignathodon Meinert, 1870 [= Rhysonotum Attems, 1952], Escaryus retusidens Attems, 1904 [= E. retusidens pallidus Folkmanová, 1956], Geophilus Leach, 1814 [= Homalarthrus Agassiz, 1846, = Esthiomenus Gistel, 1847, = Geophilus (Anadenophilus) Verhoeff, 1928], Geophilus aetnensis Verhoeff, 1928 [= G. insculptus debilis Brolemann, 1930, = G. evisensis Verhoeff, 1943, = G. henroti Manfredi, 1956, = G. aetnensis pollinensis Manfredi, 1957], G. alpinus Meinert, 1870 [= G. impressus C.L. Koch, 1847, = G. palustris C.L. Koch, 1863, = G. insculptus tauerorum Verhoeff, 1928, = G. glacialis inermis Verhoeff, 1938, = G. glacialis unguiculatus Verhoeff, 1938, = G. proximus rhenanus Verhoeff, 1895, = G. anglicanus Bagnall, 1935, = G. langkofelanus Verhoeff, 1938], G. bobolianus Verhoeff, 1928 [= G. longicornis aternanus Verhoeff, 1934], G. carpophagus Leach, 1815 [= Arthronomalus similis Newport, 1845, = G. pachymeropus Eisen & Stuxberg, 1868, = G. luridus Meinert, 1870], G. easoni Arthur, Foddai, Kettle, Lewis, Luczynski & Minelli 2001 [= Arthronomalus crassicornis Parfitt, 1866], G. electricus (Linnaeus, 1758) [= Scolopendra phosphorica Fourcroy, 1785], G. flavus (De Geer, 1778) [= G. longicornis trisulcus Silvestri, 1895, = G. longicornis glaber Verhoeff, 1928, = G. pygmaeus styricus Verhoeff, 1895, = G. longicornis pseudotruncorum Verhoeff, 1896, = G. longicornis styricorum Verhoeff, 1934, = G. carnicus praedator Verhoeff, 1937, = Pachymerium flavum Folkmanová, 1949, = G. osquidatum  porosus Dobroruka, 1957, = Schizotaenia ornata Folkmanová & Dobroruka, 1960], G. fucorum Brölemann, 1909 [= G. longicornis taorminensis Verhoeff, 1928, = G. ruinarum Verhoeff, 1931, = Pachymerium dragani Capuse, 1975], G. gavoyi Chalande, 1910 [= G. gavoyi elongatus Chalande, 1910], G. proximus C.L. Koch, 1847 [= G. ganonotus Attems, 1901, = G. eremophilus Lignau, 1933], G. pygmaeus Latzel, 1880 [= G. cispadanus Silvestri, 1896, = G. carnicus Verhoeff, 1928], G. pyrenaicus Chalande, 1909 [= G. pyrenaicus elongatus Chalande, 1909], G. seurati Brolemann, 1924 [= G. litorivagus Verhoeff, 1943], Gnathoribautia Brölemann, 1909 [= Turkomerium Chamberlin, 1952], Haplophilus Cook, 1896 [= Bothrohaplophilus Verhoeff, 1908, = Nesoporogaster Verhoeff, 1924], Haplophilus dimidiatus (Meinert, 1870) [= Himantarium gestri Pocock, 1890, = Ital ophilus sorattinus Verhoeff, 1951], H. excavatus (Verhoeff, 1924) [= Nesoporogaster hispanica Matic & Darabantu, 1969], H. souletinus Brölemann, 1907 [= H. souletinus lusitanus Verhoeff, 1925], H. subterraneus (Shaw, 1794) [= H. subterraneus elongatus Chalande & Ribaut, 1909], H. superbus (Meinert, 1870) [= Himantarium filum Meinert, 1870], Haploschendyla Verhoeff, 1900 [= Dalmatodyla Verhoeff, 1938, = Aporophilus Attems, 1903], Haploschendyla grantii (Pocock, 1891) [= Geophilus barbaricus Meinert, 1870, = H. europaea latzeli Demange, 1959], Henia bicarinata (Meinert, 1870) [= H. bicarinata elongata Brolemann, 1930, = H. bicarinata lapadensis Verhoeff, 1938], H. illyrica (Meinert, 1870) [= Chaetechelyne herzegowinensis Verhoeff, 1938, = H. illyrica absoloni Dobroruka, 1959], H. montana (Meinert, 1870) [= Chaetechelyne vesuviana pharyngealis Verhoeff, 1928], H. valida (Attems, 1927) [= Chaetechelyne osellai Matic & Darabantu, 1968], H. vesuviana (Newport, 1845) [= Scolopendra fusca Fourcroy, 1785, = Chaetechelyne sorattina Verhoeff, 1951], Hydroschendyla submarina (Grube, 1872) [= Arthronomalus littoralis Parfitt, 1874], Pachymerium coiffaiti Demange, 1959 [= P. ferrugineum maderianum Demange, 1959], P. ferrugineum (C.L. Koch, 1835) [= Geophilus caucasicus Attems, 1903, = P. tabacarui Capuse, 1968], Schendyla Bergsøe & Meinert, 1866 [= Astenoschendyla Brolemann, 1930, = Echinoschendyla Brölemann & Ribaut, 1912, = Microschendyla Brölemann & Ribaut, 1912, = Schizoschendyla Brölemann & Ribaut, 1912], Schendyla carniolensis Verhoeff, 1902 [= Poabius bistriatus C.L. Koch, 1847, = S. nemorensis quarnerana Verhoeff, 1937, = S. carniolensis clausensis Verhoeff, 1938, = S. carniolensis nivalis Verhoeff, 1938, = S. tesselata Verhoeff, 1943], S. nemorensis (C.L. Koch, 1837) [= S. nemorensis fountaini Turk, 1944, = Brachygeophilus sinionus Manfredi, 1953], S. tyrolensis (Meinert, 1870) [= Brachyschendyla montana prominens Ribaut & Brolemann, 1927, = S. montana herculis Verhoeff, 1938, = Brachyschendyla montana balcanica Kaczmarek, 1969, = Brachyschendyla dobrogica Matic & Darabantu, 1970], S. vizzavonae Léger & Duboscq, 1903 [= S. pellicensis Verhoeff, 1934, = S. incubationum Verhoeff, 1943], S. walachica Verhoeff, 1900 [= S. walachica rhodope nsis Kaczmarek, 1969], Stigmatogaster gracilis (Meinert, 1870) [= Himantarium laevipes C.L. Koch, 1847, = Geophilus ilicis Fabre, 1855, = S. gracilis robusta Attems, 1929, = Diadenoschisma gracile tyrrhenum Verhoeff, 1934, = D. gracile quarneranum Verhoeff, 1937, = S. gracilis aeserniana Attems, 1947], Strigamia acuminata (Leach, 1815) [ =Scolioplanes acuminatus brevidentatus Verhoeff, 1928, = Sc. acuminatus microdon Attems, 1904, = Sc. acuminatus pachypus Verhoeff, 1935, = Sc. italicus Verhoeff, 1928], S. crassipes (C.L. Koch, 1835) [= Scolioplanes variabilis carniolensis Verhoeff, 1895, = Sc. mediterraneus alsaticus Verhoeff, 1928, = Sc. mediterraneus carynthiacus Verhoeff, 1928, = Sc. crassipes longaronensis Verhoeff, 1935, = Sc. crassipes pegliensis Verhoeff, 1935, = Sc. crassipes faitanus Verhoeff, 1943], Thracophilus bulgaricus Verhoeff, 1926 [= T. beroni Matic & Darabantu, 1974], T. subterraneus Verhoeff, 1943 [= T. monoporus Attems, 1947].We also propose 14 new generic combinations: Dignathodon gracilis (Attems, 1952) [from Rhysonotum], Escaryus haasei (Sseliwanoff, 1884) [from Geophilus], Geophilus ibericus (Attems, 1952) [from Brachygeophilus], Geophilus pauciporus (Machado, 1952) [from Orinophilus], Gnathoribautia syriaca (Attems, 1903) [from Geophilus], Haplophilus excavatus (Verhoeff, 1924) [from Nesoporogaster], Haploschendyla splitensis (Verhoeff, 1938) [from Dalmatodyla], Henia duboscqui (Verhoeff, 1943) and H. ruffoi (Matic & Darabantu, 1968) [both from Chaetechelyne], Pachymerium minutum (Sseliwanoff, 1884) [from Geophilus], Schendyla capusei (Darabantu & Matic, 1969), S. hispanica (Attems, 1952) and S. verneri (Folkmanová & Dobroruka, 1960) [all from Brachyschendyla], Tuoba zograffi (Brölemann, 1900) [from Geophilus].Lectotypes have been selected for two species: Geophilus pusillus Meinert, 1870 and Himantarium mediterraneum Meinert, 1870.In seven cases we suggest to conserve currently used names over senior synonyms or homonyms: Geophilus alpinus Meinert, 1870 [over G. impressus C.L. Koch, 1847 and G. palustris C.L. Koch, 1863], Geophilus easoni Arthur, Foddai, Kettle, Lewis, Luczynski & Minelli 2001 [over Arthronomalus crassicornis Parfitt, 1866], Gnathoribautia bonensis (Meinert, 1870) [over Necrophloeophagus punctiventris Newport, 1844], Bothriogaster signata (Kessler, 1874) [over Notiphilus sanguineus C.L. Koch, 1847 and N. taeniatus C.L. Koch, 1847], Schendyla carniolensis Verhoeff, 1902 [over Poabius bistriatus C.L. Koch, 1847]. Corresponding applications have been submitted to the International Commission on Zoological nomenclature for a ruling under the Plenary Powers. 

A comprehensive dataset of the geophilid centipedes of the south-eastern Alps (Chilopoda, Geophilomorpha, Geophilidae s.l.).

Background: Centipedes of the family Geophilidae s.l. are widespread in the Holarctic, with the south-eastern part of the European Alps standing out as one of the most investigated regions. However, retrieving the published records for this taxon, even for this region, is challenging, since most of them are sparse in the specialised literature and interpreting them is hampered by the many taxonomic and nomenclatorial changes occurred in the past and recent times. New information: We assembled and released a dataset of occurrence records of the geophilid species in the south-eastern Alps, including all the published records and many other records present in unpublished catalogues of scientific collections. For each record, we integrated information from all the sources about: locality, date of collection, the taxonomic identifications, number and sex of individuals and available sequences of molecular markers. For all the records, we estimated geographic coordinates of the locality, when not originally provided, based on the information available. We also estimated the accuracy of the position.The dataset includes 3293 records referred to 39 species, obtained since the first half of the 19th century and up to 2022; 52% of these records have been released publicly for the first time in the dataset here described.

An updated synthesis of the Geophilomorpha (Chilopoda) of Asian Russia.

A comprehensive overview of the state of knowledge on the ChilopodaGeophilomorpha of the Asian part of Russia is presented, based on the critical revision of all published morphological descriptions and all geographical records. Revised diagnoses for all the 38 nominal species so far reported from Asian Russia are given, with comments on their validity. Among them a total of 18 species are recorded only from this region and many of them from a single locality only. The species belong to Geophilidae s. l., (in the genera Arctogeophilus, Geophilus, Pachymerium, and Strigamia), Schendylidae (Escaryus), and Mecistocephalidae (Agnostrup, Arrup, and Tygarrup). At least two species have been introduced, namely Geophilusflavus and Tygarrupjavanicus. The history of studies on the Geophilomorpha in the Asian part of Russia are also summarized.

Alpine salamanders at risk? The current status of an emerging fungal pathogen.

Amphibians globally suffer from emerging infectious diseases like chytridiomycosis caused by the continuously spreading chytrid fungi. One is Batrachochytrium salamandrivorans (Bsal) and its disease ‒ the 'salamander plague' ‒ which is lethal to several caudate taxa. Recently introduced into Western Europe, long distance dispersal of Bsal, likely through human mediation, has been reported. Herein we study if Alpine salamanders (Salamandra atra and S. lanzai) are yet affected by the salamander plague in the wild. Members of the genus Salamandra are highly susceptible to Bsal leading to the lethal disease. Moreover, ecological modelling has shown that the Alps and Dinarides, where Alpine salamanders occur, are generally suitable for Bsal. We analysed skin swabs of 818 individuals of Alpine salamanders and syntopic amphibians at 40 sites between 2017 to 2022. Further, we compiled those with published data from 319 individuals from 13 sites concluding that Bsal infections were not detected. Our results suggest that the salamander plague so far is absent from the geographic ranges of Alpine salamanders. That means that there is still a chance to timely implement surveillance strategies. Among others, we recommend prevention measures, citizen science approaches, and ex situ conservation breeding of endemic salamandrid lineages.

Morphology of a neglected large-sized species of Geophilus from Iran (Chilopoda: Geophilidae).

Geophilus gigas was described briefly by Attems in 1951 from two sites in Iran, but neither reported nor cited subsequently. The original description is inadequate to understand the full morphology and the taxonomic position of the species. Thus it is here redescribed and figured in detail based on the examination of a syntype and newly collected specimens. Furthermore it is compared with the most similar species of the genus Geophilus. G. gigas can be reliably distinguished from other related species for the maximum body size (more than 6 cm), the arrangement and extent of the sternal pore-fields (with additional groups of pores), the coxal pores number and distribution (tens, scattered), and the size of the ultimate legs (swollen in both sexes).

Diversity and biogeography of Israeli geophilomorph centipedes (Chilopoda: Geophilomorpha).

The fauna of Chilopoda Geophilomorpha of Israel has been analyzed after examining 128 new specimens from 35 localities, reinterpreting all published data including 103 records, and relating occurrence of species with major climatic parameters. A key to identification has been compiled. A total of 17 species are distinguished, of which three are reported from Israel for the first time, while five are documented by published records only. The following new synonymies are proposed and discussed: Dignathodon pachypus Verhoeff, 1943 = Dignathodon microcephalus (Lucas, 1846); Geophilus flavidus noduliger Verhoeff, 1925 = Clinopodes escherichii (Verhoeff, 1896); Pachymerium ferrugineum vosseleri Verhoeff, 1902 = P. ferrugineum (Koch, 1835). Of all the species, Bothriogaster signata and Pachymerium ferrugineum are widespread in the country, while other species occupy different climatic zones, from desert to more humid and montane.

The centipede fauna (Chilopoda) of the island of Cyprus, with one new lithobiomorph species.

The centipede (Chilopoda) fauna of Cyprus, which was almost unknown, has been analysed by examining more than 1,800 specimens sampled from 185 sites, besides revising critically the few published data. A total of 26 species are listed and discussed (1 Scutigeromorpha, 9 Lithobiomorpha, 3 Scolopendromorpha, 13 Geophilomorpha), 21 of which are new to the island, i.e. Scutigera coleoptrata (Linnaeus, 1758), Lithobius (Ezembius) parvicornis (Porat, 1893), L. (E.) pamukkalensis Matic, 1980, L. (E.) zeylanus (Chamberlin, 1952), L. (Lithobius) carinatus L. Koch, 1862, L. (L.) erythrocephalus C.L. Koch, 1847, L. (Lithobius?) anderssoni n. sp., L. (Monotarsobius) ferganensis Trotzina, 1880, Cryptops (Cryptops) kosswigi (Chamberlin, 1952), C. (C.) cf. trisulcatus Brölemann, 1902, Dignathodon microcephalus (Lucas, 1846), Henia (Meinertia) bicarinata (Meinert, 1870), Geophilus cf. alpinus Meinert, 1870, G cf. carpophagus Leach, 1815, Pachymerium ferrugineum (C.L. Koch, 1835), Schizotaenia sp., Stenotaenia naxia (Verhoeff, 1901), Thracophilus cilicius Attems, 1947, Nannophilus eximius (Meinert, 1870) and Schendyla cf. nemorensis (C.L. Koch, 1837), and another unidentified species of Schendylidae . As far as known, Lithobius anderssoni n. sp. is endemic to the island. Geographic distribution in Cyprus and ecological notes are given for each species. Taxonomic remarks are given for some species.

Soil centipedes (Chilopoda, Geophilomorpha) in the Val Camonica forests (Southern Alps): species composition and richness.

Soil centipedes (Chilopoda, Geophilomorpha) are a widespread group of predators in the forest soils of the European Alps. While in the eastern and western parts of the Southern Prealps, larger efforts were devoted to sample and study the geophilomorph fauna, little is known about species richness and composition of geophilomorph communities in the central part of Southern Prealps. In this work, five sites located in the Val Camonica were surveyed by hand searching, between November 2021 and July 2022 and their species richness was estimated applying non-parametrical statistical methods (Chao-1 and Abundance-based Coverage Estimator) to account for incomplete detection. A total of 18 species were found amongst the five sites. A maximum of 12 species were recorded in each single site, while estimates suggest that another 1-3 species were likely undetected. Species composition were found highly variable also between sites with similar species richness.

A new giant species of Clinopodes from the Alborz Mountains, at the eastern edge of the genus range (Chilopoda: Geophilidae).

A new species of Clinopodes C. L. Koch, 1847, Clinopodes azadi n. sp., is described and illustrated, based on two specimens from Alborz Mountains, Iran. The new species lives in the easternmost part of the Western Palearctic range of the genus and is outstanding in comparison to all other known Clinopodes species, being the largest (surpassing 11 cm in length) and with the highest number of body segments (89-91 leg-bearing segments). C. azadi n. sp. differs from the other Clinopodes species also in the coxosternite of first maxillae (with a distinct mid-longitudinal sutural sulcus) and in the coxal pores (numerous, and sparse on the entire surface of the coxopleura).

Morphology and distribution of the Middle Asian centipede genus Krateraspis Lignau, 1929 (Chilopoda, Geophilomorpha, Mecistocephalidae).

A comprehensive redescription of the poorly known mecistocephalid genus Krateraspis Lignau, 1929 and its two species is provided, based on the examination of type material and new specimens, as well as the critical evaluation of all published information. Krateraspis is confirmed differing from all other Mecistocephalidae especially for a peculiar pattern of areolation and setation of the clypeus. Records from 24 localities indicate that Krateraspis is limited to a narrow area of Middle Asia, from the Western Tian-Shan to the western offshoots of Pamir Mountains. Two species are morphologically distinguishable: K.meinerti (Sseliwanoff, 1881) and K.sselivanovi Titova, 1975. They differ mainly in details of the clypeus and maxillae, in the pattern of forcipular denticles, and in the number of legs. Tygarrupasiaticus Verhoeff, 1930 is confirmed as a junior synonym of K.meinerti, and a lectotype is designated for the former.

Geophilomorpha from Alborz Mountains and a checklist of Chilopoda from Iran.

This study presents an updated checklist of centipedes (Chilopoda) from Iran based on a review of the literature and recently examined specimens of Geophilomorpha collected from many sites in the Alborz Mountains. The centipedes of Iran comprise 48 species, of which 17 species have so far only been reported from Iran. Many species are known from a single locality, and only from specimens of a single sex. Species of the geophilomorph genera Clinopodes and Strigamia are here reported from Iran for the first time. The checklist includes all published geographical distribution data from Iran and a comparison with the centipede fauna of neighboring countries using the Jaccard index. The Iranian centipede fauna is still scantily known and further investigations are expected to contribute substantially, especially because of the diverse physiography of the country.

A new soil centipede from South-East Asia with a unique arrangement of ventral glands, and a revised synopsis of Gonibregmatidae (Chilopoda, Geophilomorpha).

A new gonibregmatid centipede, Vinaphilusunicus gen. n., sp. n., is described based on two females from a single location in northern Vietnam. The new genus and species are distinguished mainly by the arrangement of the ventral pore-fields, which is unique among all Chilopoda. A critically revised synopsis of the Gonibregmatidae is also given. In particular, three species are provisionally recognized in Himatosoma Pocock, 1891: H.bidivisum Silvestri, 1919, H.porosum Pocock, 1891 (= H.typicumtridivisum Silvestri, 1919, syn. n.), and H.typicum Pocock, 1891. The genera Dschangelophilus Verhoeff, 1937 and Tweediphilus Verhoeff, 1937, with their species D.coloratus Verhoeff, 1937 and T.malaccanus Verhoeff, 1937, are moved to the Gonibregmatidae, whereas Geoporophilusaporus Attems, 1930 is moved to the Oryidae as Orphnaeusaporus (Attems, 1930), comb. n.

Revision of the Neotropical species of Aphilodontinae (Geophilomorpha, Geophilidae), with eight new species and a first phylogenetic analysis of the subfamily.

Aphilodontinae are a well distinct but poorly known group of soil centipedes (Chilopoda: Geophilomorpha: Geophilidae) inhabiting two disjunct areas in South America and South Africa respectively. A comprehensive revision of the Neotropical species is presented based on the examination of >150 specimens, the phylogenetic analysis of the entire subfamily based on 47 morphological characters, and the description of eight new species from Southeastern Brazil. The Aphilodontinae were confirmed a monophyletic clade within Geophilidae and closely related to Geoperingueyia, with which they share synapomorphic characters in the labrum, second maxillae and forcipules. Particular groups of clypeal setae, inconspicuous lateral parts of labrum and coalescent forcipular throchanteroprefemur and femur were recovered as synapomorphies of the Aphilodontinae. Four monophyletic genera are recognized within the subfamily Aphilodontinae Silvestri, 1909 (1908) (= Brasilophilidae Verhoeff, 1908; = Mecistaucheninae Verhoeff, 1925 n. syn.), three exclusively Neotropical and one exclusively South African. The Neotropical genera are: Aphilodon Silvestri, 1898 (= Mecistauchenus Brölemann, 1907 n. syn.; = Brasilophilus Verhoeff, 1908 n. syn.), with ten species (including A. caboclos n. sp., A. indespectus n. sp., A. meganae n. sp., A. pereirai n. sp. and A. silvestrii n. sp.), characterized by a small terminal spine on the ultimate legs, in both males and females; Mecophilus Silvestri, 1909, with two species (including M. tupiniquim n. sp.), characterized by elongated forcipular segment and first leg-bearing segment; Mairata n. gen., with two species (M. butantan n. sp. and M. itatiaiensis n. sp.), characterized by the partial reduction of the distal article of the second maxillary telopodite and a particular shape of ultimate legs, in both males and females. The South African genus, Philacroterium Attems, 1926, comprises eleven known species (including P. weberi (Silvestri, 1909) n. comb., P. caffrarium (Verhoeff, 1937) n. comb., P. porosum (Verhoeff, 1937) n. comb., P. brevipes (Verhoeff, 1938) n. comb., P. bidentatum (Lawrence, 1955) n. comb., P. longipes (Lawrence, 1955) n. comb., P. macronyx (Lawrence, 1955) n. comb., P. maritimum (Lawrence, 1963) n. comb., P. pauciporum (Lawrence, 1963) n. comb. and P. transvaalicum (Lawrence, 1963) n. comb.) and is characterized by vestigial lateral parts of the labrum and the complete suppression of the distal article of the telopodites of the second maxillae. Identification keys are provided for the Aphilodontinae genera and for the species of Aphilodon, Mecophilus and Mairata n. gen. The known occurrence of Aphilodon in Brazil is extended from the states of São Paulo and Mato Grosso to the states of Minas Gerais and Rio de Janeiro, that of Mecophilus from the state of Paraná to the state of São Paulo.

Geophilus pygmaeus (Chilopoda: Geophilidae): clarifying morphology, variation and geographic distribution.

The European centipede Geophilus pygmaeus (Chilopoda: Geophilidae) has been known since 1880. It has been repeatedly reported since then and its validity has never been questioned. Nevertheless, a complete description of the species and an unambiguous diagnosis are still lacking. Based on the examination of one of the syntypes and > 110 specimens from 28 other localities, we provide here a full description and illustration of the morphology of G. pygmaeus and a revised and updated overview of the geographic range of the species. G. pygmaeus is less than 2 cm long, has 41-47 pairs of legs and differs from all other European species of Geophilus in a combination of characters including the shape of the forcipular apparatus and a unique arrangement of the coxal pores. G. pygmaeus is distributed through the Southern Limestone Alps, between the Bergamasque Prealps in the West and the Slovene Prealps in the East, and in the northernmost Dinarides. Published records from other European areas are actually erroneous or uncertain.

The first geophilid centipedes from Malesia: a new genus with two new species from Sumatra (Chilopoda, Geophilidae).

A new genus Sundageophilus is here described for two new species of geophilid centipedes (Chilopoda: Geophilidae) from Sumatra, Indonesia. Both Sundageophilus bidentatus sp. n. and Sundageophilus poriger sp. n. feature a minute body size (less than 1 cm long with 31-35 pairs of legs), a similar structure of the maxillae, elongated forcipules, and few coxal organs. Sundageophilus bidentatus is unique among geophilids because the ultimate article of the forcipule is armed with two conspicuous denticles, one dorsal to the other, instead of a single one or none. The two species of Sundageophilus are the first genuine Geophilidae ever found in Malesia, and among the very few representatives of this family in the entire south-eastern Asia.

At the end of the rope: Geophilushadesi sp. n. - the world's deepest cave-dwelling centipede (Chilopoda, Geophilomorpha, Geophilidae).

A new geophilomorph centipede, Geophilushadesi sp. n., is described from caves in the Velebit Mountain, central Croatia. Together with Geophiluspersephones Foddai & Minelli, 1999, described from Pierre Saint-Martin cave in France, they are the only two remarkably troglomorphic geophilomorphs hitherto known. The new species apparently belongs to a group of Geophilus species inhabiting mainly Western and Southern Europe, with a uniquely modified pretarsus in the second maxillae. Geophilushadesi sp. n. shows unusual traits, some of which commonly found in troglobitic arthropods, including exceptionally elongated antennae, trunk segments and leg claws. The species is described upon specimens found in two caves at a depth below -250 m. Another two specimens apparently belonging to the same species have been recorded in another deep vertical cave at -980 m and -1100 m. The latter represents the world's deepest record of Chilopoda as a whole.

Molecular-based estimate of species number, phylogenetic relationships and divergence times for the genus Stenotaenia (Chilopoda, Geophilomorpha) in the Italian region.

Stenotaenia is one of the largest and most widespread genera of geophilid centipedes in the Western Palearctic, with a very uniform morphology and about fifteen species provisionally recognized. For a better understanding of Stenotaenia species-level taxonomy, we have explored the possibility of using molecular data. As a preliminary assay, we sampled twelve populations, mainly from the Italian region, and analyzed partial sequences of the two genes COI and 28S. We employed a DNA-barcoding approach, complemented by a phylogenetic analysis coupled with divergence time estimation. Assuming a barcoding gap of 10-16% K2P pairwise distances, we found evidence for the presence of at least six Stenotaenia species in the Italian region, which started diverging about 50 million years ago, only partially matching with previously recognized species. We found that small-sized oligopodous species belong to a single clade that originated about 33 million years ago, and obtained some preliminary evidence of the related genus Tuoba being nested within Stenotaenia.