The millipede genera Amblyiulus Silvestri, 1896 and Syrioiulus Verhoeff, 1914 in the Caucasus, with notes on their distributions (Diplopoda, Julida, Julidae).

In the Caucasus, the genera Amblyiulus Silvestri, 1896 and Syrioiulus Verhoeff, 1914 are shown to include two and four species, respectively: Amblyiulus georgicus Lohmander, 1932, from Georgia and Armenia, A. hirtus sp. nov., from Azerbaijan and Dagestan, Russia, Syrioiulus adsharicus (Lohmander, 1936), from Georgia, S. continentalis (Attems, 1903), from Azerbaijan and Iran, S. taliscius (Attems, 1927), from Azerbaijan, and S. armeniacus sp. nov., from Armenia. All these six species are described, illustrated, and keyed, and their distributions are mapped and discussed, based on the literature data and abundant new samples.

Macedomeris, a new monotypic doderiinine genus (Diplopoda, Glomerida) from a cave in North Macedonia, Balkan Peninsula.

Here we describe a new monotypic glomerid genus, Macedomeris gen. nov., with Macedomeris ivoi sp. nov. (a presumed troglobiont) as its type species. The new genus clearly differs from all other genera within the order Glomerida by the combination of several morphological characters: the presence of a characteristic deep lateral pit ("Ohrgrube") on both sides of the thoracic shield, the absence of striking ornamentation on tergites, the fusion of tergite 11 with the anal shield, and the lack of a large medial hump. In addition, Macedomeris gen. nov. differs from other members of the Glomerida by its general appearance, the presence of 2 or 3 vestigial ommatidia and a very wide syncoxite of a 2-segmented leg-pair 17 in the male. Notes on its ecology, troglomorphic features, and relationships with similar members of the tribe Doderiini are also given.

Geophilus serbicus sp. nov., a new species from the Balkan Peninsula (Chilopoda: Geophilomorpha: Geophilidae).

Geophilus serbicus sp. nov., a new epigeic centipede species, is described and illustrated based on specimens collected from the Stara Planina Mountains (the Balkan Mountain Range), Eastern Serbia, Balkan Peninsula. Considerations on the taxonomic relationships with some similar Geophilus species are briefly presented, and the distribution of the new species is mapped.

Review of the genus Typhloiulus Latzel, 1884 in the Dinaric region, with a description of four new species and the first description of the male of Typhloiulus insularis Strasser, 1938 (Diplopoda: Julida: Julidae).

A review of the genus Typhloiulus Latzel, 1884 (Julida: Julidae) in the Dinaric region is given. Previously published and new faunistic records are provided for each species, together with notes for some of them. Four new species are described: T. clavatus sp. nov., T. gracilis sp. nov., T. opisthonodus sp. nov., and T. parvulus sp. nov., all from caves in Dalmatia and Herzegovina. In addition, we provide the first description of the male of T. insularis Strasser, 1938. Some taxonomic relationships among Typhloiulus species and biogeographical aspects (with distribution maps) of the genus are discussed.

From the depths: Heterocaucaseuma deprofundum sp. nov., the world's deepest-occurring millipede (Diplopoda, Chordeumatida, Anthroleucosomatidae) from caves in the western Caucasus.

We describe a new anthroleucosomatid millipede, Heterocaucaseuma deprofundum sp. nov., found in the second and third deepest caves of our planet, Krubera-Voronja and Sarma, in Abkhazia, western Caucasus. A brief discussion on its taxonomy and relationships within the Caucaseuma complex and genus Heterocaucaseuma is provided, together with new faunistic records on other three species of Heterocaucaseuma. Notes on the distribution and troglomorphy of the genus Heterocaucaseuma are added. Ecological information and data on the subterranean community of the Krubera-Voronja Cave are updated. We also discuss the vertical distribution of the new species in caves and the presence of an ectoparasitic fungi of the order Laboulbeniales. This new species is currently the world's deepest-occurring millipede, also representing one of the largest known chordeumatidans.

Dalmatosomatidae, a new monotypic family, and Dalmatosoma agaricum gen. et sp. nov. (Diplopoda: Chordeumatida: Craspedosomatidea) from Croatia, Balkan Peninsula.

A new millipede species, Dalmatosoma agaricum gen. et sp. nov., is described from a cave in Dalmatia (Croatia). The new taxon is clearly unique among chordeumatidans in both anterior and posterior gonopod structure and shows some noteworthy habitus features. We therefore here establish a new monotypic family, viz., Dalmatosomatidae, for the given taxon and place it within the suborder Craspedosomatidea. The new family is briefly discussed and compared with all other chordeumatidans.

Revision of the genus Hylopachyiulus Attems, 1904, with a description of a new species from Croatia (Diplopoda, Julida, Julidae).

A revision of the julid genus Hylopachyiulus Attems, 1904 is given, and lectotypes and paralectotypes are respectively designated for the species H. corylorum (Verhoeff, 1907) and H. pygmaeus (Attems, 1904). We confirm the validity of H. pygmaeus and synonymize H. likanus Attems, 1926 syn. nov. under H. corylorum which is resurrected. Additonally, a third species is described from Croatia-H. ocellatus sp. nov.. Considerations on the taxonomic relationships in the tribe Pachyiulini and within the genus Hylopachyiulus are presented, and the distribution of its species is illustrated and discussed.

Chemical Ecology of Cave-Dwelling Millipedes: Defensive Secretions of the Typhloiulini (Diplopoda, Julida, Julidae).

Cave animals live under highly constant ecological conditions and in permanent darkness, and many evolutionary adaptations of cave-dwellers have been triggered by their specific environment. A similar "cave effect" leading to pronounced chemical interactions under such conditions may be assumed, but the chemoecology of troglobionts is mostly unknown. We investigated the defensive chemistry of a largely cave-dwelling julid group, the controversial tribe "Typhloiulini", and we included some cave-dwelling and some endogean representatives. While chemical defense in juliform diplopods is known to be highly uniform, and mainly based on methyl- and methoxy-substituted benzoquinones, the defensive secretions of typhloiulines contained ethyl-benzoquinones and related compounds. Interestingly, ethyl-benzoquinones were found in some, but not all cave-dwelling typhloiulines, and some non-cave dwellers also contained these compounds. On the other hand, ethyl-benzoquinones were not detected in troglobiont nor in endogean typhloiuline outgroups. In order to explain the taxonomic pattern of ethyl-benzoquinone occurrence, and to unravel whether a cave-effect triggered ethyl-benzoquinone evolution, we classed the "Typhloiulini" investigated here within a phylogenetic framework of julid taxa, and traced the evolutionary history of ethyl-benzoquinones in typhloiulines in relation to cave-dwelling. The results indicated a cave-independent evolution of ethyl-substituted benzoquinones, indicating the absence of a "cave effect" on the secretions of troglobiont Typhloiulini. Ethyl-benzoquinones probably evolved early in an epi- or endogean ancestor of a clade including several, but not all Typhloiulus (basically comprising a taxonomic entity known as "Typhloiulus sensu stricto") and Serboiulus. Ethyl-benzoquinones are proposed as novel and valuable chemical characters for julid systematics.

The first hydrophilous cave-dwelling millipede from Serbia-Typhloiulus balcanicus sp. nov. (Diplopoda, Julida, Julidae).

As the fourth hydrophilous species of the genus Typhloiulus Latzel, 1884, the new species T. balcanicus Antic sp. nov. is described from two caves in the Stara Planina Mountains of Serbia. A brief discussion of the relationships between the new species and congeners with modified and unmodified mouthparts is presented, as well as short notes on the habitat of the new species and comments on other semiaquatic millipedes with modified mouthparts.

A new cave-dwelling species of the genus Mesoiulus Berlese, 1886 (Diplopoda, Julida, Julidae) from Turkey.

A new cave-dwelling species of the poorly defined genus Mesoiulus Berlese, 1886, M. taurus sp. nov., is described from the Zindan Cave, Isparta, Western Taurides, Turkey. A brief discussion of the relationship between M. taurus sp. nov. and the most similar species (M. ciliciensis Strasser, 1975) is presented, as well as a map showing the distribution of the genus Mesoiulus in Turkey.

The Caucasus as a major hotspot of biodiversity: Evidence from the millipede family Anthroleucosomatidae (Diplopoda, Chordeumatida).

Fifteen new genera and 36 new species of the diplopod family Anthroleucosomatidae Verhoeff, 1899 are described from the Caucasus, one of the world's biodiversity hotspots: Golovatchosoma bacillichaetum gen. et sp. nov., G. trichochaetum sp. nov., Alpinella waltheri gen. et sp. nov., Brachychaetosoma turbanovi gen. et sp. nov., Caucaseuma elephantum sp. nov., C. fanagoriyskaya sp. nov., C. glabroscutum sp. nov., C. kelasuri sp. nov., C. minellii sp. nov., C. variabile sp. nov., Caucasominorus billi gen. et sp. nov., C. polylobatus sp. nov., Georgiosoma bicornutum gen. et sp. nov., Heterocaucaseuma feminaepectorum gen. et sp. nov., H. longicorne sp. nov., Paranotosoma attemsi gen. et sp. nov., P. cordatum sp. nov., P. subrotundatum sp. nov., Dentatosoma denticulatum gen. et sp. nov., D. magnum sp. nov., D. zeraboseli sp. nov., Enghoffiella insolita gen. et sp. nov., Metamastigophorophyllon hamatum sp. nov., M. lamellohirsutum sp. nov., M. torsivum sp. nov., Acanthophorella barjadzei gen. et sp. nov., A. chegemi sp. nov., A. irystoni sp. nov., Cryptacanthophorella manubriata gen. et sp. nov., Flagellophorella hoffmani gen. et sp. nov., Pseudoflagellophorella eskovi gen. et sp. nov., P. mirabilis sp. nov., P. papilioformis sp. nov., Herculina oligosagittae gen. et sp. nov., H. polysagittae sp. nov., Vegrandosoma tabacarui gen. et sp. nov. All anthroleucosomatids from the Caucasus, both new and previously described, are classified in eight endemic complexes, arranged as follows in alphabetic order: the Alloiopus complex, the Caucaseuma complex, the Dentatosoma complex, the Enghoffiella complex, the Flagellophorella complex, the Herculina complex, the Ratcheuma complex and the Vegrandosoma complex. The genus Metamastigophorophyllon Ceuca, 1976 (the type-species: Mastigophorophyllon giljarovi Lang, 1959, adequately redescribed from new material), is a new subjective senior synonym of Persedicus Mauriès, 1982, syn. nov., being also newly transferred from the family Mastigophorophyllidae to the family Anthroleucosomatidae, comb. nov. The following two new combinations are also proposed: Heterocaucaseuma mauriesi (Golovatch and Makarov, 2011), comb. nov. ex Anamastigona Silvestri, 1898; and Metamastigophorophyllon martensi (Mauriès, 1982), comb. nov. ex Persedicus Mauriès, 1982. An identification key to and distribution maps for all known species of Caucasian Anthroleucosomatidae are provided, as well as brief remarks on distributions and on cave-dwelling taxa.

"Quinone Millipedes" Reconsidered: Evidence for a Mosaic-Like Taxonomic Distribution of Phenol-Based Secretions across the Julidae.

The defensive chemistry of juliformian millipedes is characterized mainly by benzoquinones ("quinone millipedes"), whereas the secretions of the putative close outgroup Callipodida are considered to be exclusively phenolic. We conducted a chemical screening of julid secretions for phenolic content. Most species from tribes Cylindroiulini (15 species examined), Brachyiulini (5 species examined), Leptoiulini (15 species examined), Uncigerini (2 species examined), Pachyiulini (3 species examined), and Ommatoiulini (2 species examined) had non-phenolic, in most cases exclusively benzoquinonic secretions. In contrast, tribes Cylindroiulini, Brachyiulini, and Leptoiulini also contained representatives with predominantly phenol-based exudates. In detail, p-cresol was a major compound in the secretions of the cylindroiulines Styrioiulus pelidnus and S. styricus (p-cresol content 93 %) and an undetermined Cylindroiulus species (p-cresol content 51 %), in the brachyiulines Brachyiulus lusitanus (p-cresol content 21 %) and Megaphyllum fagorum (p-cresol content 92 %), as well as in an undescribed Typhloiulus species (p-cresol content 32 %, Leptoiulini). In all species, p-cresol was accompanied by small amounts of phenol. The secretion of M. fagorum was exclusively phenolic, whereas phenols were accompanied by benzoquinones in all other species. This is the first incidence of clearly phenol-dominated secretions in the Julidae. We hypothesize a shared biosynthetic route to phenols and benzoquinones, with benzoquinones being produced from phenolic precursors. The patchy taxonomic distribution of phenols documented herein supports multiple independent regression events in a common pathway of benzoquinone synthesis rather than multiple independent incidences of phenol biosynthesis.

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.

A new species of the family Heterolatzeliidae from the Balkan Peninsula (Diplopoda, Chordeumatida).

A new species of Heterolatzelia Verhoeff, 1897, H. karlstrasseri sp. n., is described from the Datlo Cave in east Herzegovina. A brief discussion about the relationship between H. karlstrasseri sp. n. and H. durmitorensis Gulicka, 1968 and a distribution map of the family Heterolatzeliidae are presented. 

Three new monotypic genera of Trichopolydesmidae from Croatia, Balkan Peninsula (Diplopoda, Polydesmida).

Three new monotypic genera are described from underground habitats in Croatia: Balkanodesmus biokovensis gen. n., sp. n., Solentanodesmus insularis gen. n., sp. n. and Velebitodesmus cavernicolus gen. n., sp. n. In all three new genera the prefemoral part of the gonopods is orientated transversely to the main body axis, and all three new genera possess numerous metatergal setae arranged in a few transverse irregular rows, a condition observed in several European trichopolydesmids. These three new genera are very close to the genus Verhoeffodesmus. A distribution map and a brief discussion about the relationship with congeners are provided.

Chemical defense in millipedes (Myriapoda, Diplopoda): do representatives of the family Blaniulidae belong to the 'quinone' clade?

The defensive secretions of two blaniulid millipedes, Nopoiulus kochii and Cibiniulus phlepsii, were characterized by GC-FID and GC/MS analyses, which showed the presence of a complex mixture of benzoquinones, hydroquinones, and oleates. Altogether, 13 compounds were identified. The major compound in the secretions of both analyzed species was 2-methyl-1,4-benzoquinone (toluquinone). The second major constituent in the N. kochii secretion was 2-methyl-3,4-(methylenedioxy)phenol, while in that of C. phlepsii, it was 2-methoxy-3-methyl-1,4-benzoquinone. The defensive secretion of N. kochii also showed a high content of hydroquinones (13.5%) in comparison to that of C. phlepsii (0.8%). Hexyl oleate and octyl oleate were detected for the first time in defensive millipede fluids. The chemical composition of the defensive secretions supports the chemotaxonomic position of the family Blaniulidae in the 'quinone' millipede clade.

Chemistry of the sternal gland secretion of the Mediterranean centipede Himantarium gabrielis (Linnaeus, 1767) (Chilopoda: Geophilomorpha: Himantariidae).

The geophilomorph centipede, Himantarium gabrielis, when disturbed, discharges a viscous and proteinaceous secretion from the sternal glands. This exudate was found by gas chromatography-mass spectrometry, liquid chromatography-high resolution mass spectrometry, liquid chromatography-mass spectrometry-mass spectrometry and NMR analyses to be composed of hydrogen cyanide, benzaldehyde, benzoyl nitrile, benzyl nitrile, mandelonitrile, mandelonitrile benzoate, 3,7,6O-trimethylguanine (himantarine), farnesyl 2,3-dihydrofarnesoate and farnesyl farnesoate. This is the first report on the presence of benzyl nitrile and mandelonitrile benzoate in secreted substances from centipedes. Farnesyl 2,3-dihydrofarnesoate is a new compound, while himantarine and farnesyl farnesoate were not known as natural products. A post-secretion release of hydrogen cyanide by reaction of mandelonitrile and benzoyl nitrile was observed by NMR, and hydrogen cyanide signals were completely assigned. In addition, a protein component of the secretion was analysed by electrophoresis which revealed the presence of a major 55 kDa protein. Analyses of the defensive exudates of other geophilomorph families should produce further chemical surprises.