Catamicrophyllum beroni sp. nov.the first European record of the genus Catamicrophyllum Verhoeff, 1901 (Diplopoda: Julida: Julidae).

A new species of Catamicrophyllum Verhoeff, 1901, Catamicrophyllum beroni sp. nov., is described from Rila Mts, southwest Bulgaria. The new species is compared to its most similar congener, Catamicrophyllum caucasicum (Attems, 1901) known from the Caucasus and Eastern Anatolia, including descriptive notes and scanning electron micrographs of the latter. The remarkably disjunct distribution areas of the two species are briefly discussed in the context of similar distribution patterns in other members of Julidae. In addition, some ecological notes and conservation issues in connection with the climatic changes occurring in the area of the type locality of C. beroni sp. nov. are presented.

A review of the genus Micropachyiulus Verhoeff, 1899 and description of the related Armeniopachyiulus gen. nov. (Diplopoda: Julida: Julidae: Pachyiulini).

The European millipede genus Micropachyiulus Verhoeff, 1899 is revised based on type and non-type material, and on the available literature. The genus Hylopachyiulus Attems, 1904 is established as a junior subjective synonym of Micropachyiulus, syn. nov. Micropachyiulus ocellatus (Antic & Akkari, 2018) is thus listed as a new combination, comb. nov. Two new species-Micropachyiulus caucasicus sp. nov. and M. filiformis sp. nov.-are described from Russia and Romania, respectively. In addition, a presumably closely related monospecific genus, Armeniopachyiulus gen. nov., and its sole species A. pokr gen. et sp. nov. are described from Armenia. A brief discussion on the systematics and distribution of the studied taxa is presented.

A review of the cavernicolous Trichopolydesmidae (Diplopoda, Polydesmida) from the Carpathian-Balkan arch and the Rhodope Mountains, with descriptions of two new genera and three new species.

All cavernicolous species of the millipede family Trichopolydesmidae from the Carpathian-Balkan arch and the Rhodope Mountains have been reviewed. At present the family has been shown to comprise five or six genera with eight or nine species. Two new genera have been described, viz., Balkanodesminus gen. nov., with two new species: B.dentatoides sp. nov. and B.serbicus sp. nov., from Bulgaria and Serbia, respectively, and the monospecific Rhodopodesmus gen. nov., with R.niveus sp. nov., from Bulgaria. Two new combinations and one new status have been proposed: Balkanodesminusbulgaricus (Strasser, 1962) comb. nov. ex Bacillidesmusbulgaricus Strasser, 1962 and Balkanodesminusdentatus (Strasser, 1966a) comb. nov., stat. nov. ex Bacillidesmusbulgaricusdentatus Strasser, 1966a. All genera and species are diagnosed with the inclusion of the most relevant remarks for each of them. Old museum types are checked for Bacillidesmusfiliformis (Latzel, 1884) with lectotype designation, as well as for Trichopolydesmuseremitis Verhoeff, 1898. An identification key to all six genera and a distribution map for the eight species are provided, as well as brief remarks and general considerations on the family Trichopolydesmidae.

A new millipede genus and species of the tribe Pachyiulini from the Caucasus (Diplopoda, Julida, Julidae).

A new genus and species of the millipede tribe Pachyiulini, Bellatoiulusgolovatchi gen. et sp. nov., is described from the Lesser Caucasus, Azerbaijan. Cybertypes of the new species are created from the physical holotype male and from a paratype female. The distribution and ecological features of the new species, and the position of the new genus within Pachyiulini are discussed.

The millipede tribe Brachyiulini in the Caucasus (Diplopoda, Julida, Julidae).

The diplopod tribe Brachyiulini is represented in the fauna of the Caucasus by eight genera and 32 species, of which one genus and 14 species are described as new: Colchiobrachyiulusmontanus Vagalinski, sp. nov., Iraniulustricornis Vagalinski, sp. nov., Omobrachyiulusarmatus Vagalinski, sp. nov., O.fasciatus Vagalinski, sp. nov., O.faxifer Vagalinski, sp. nov., O.kvavadzei Vagalinski, sp. nov., O.lazanyiae Vagalinski, sp. nov., O.ponticus Vagalinski, sp. nov., O.pristis Vagalinski, sp. nov., O.trochiloides Vagalinski, sp. nov., O.unugulis Vagalinski, sp. nov., O.zuevi Vagalinski, sp. nov., Svaniulusryvkini Vagalinski, gen. nov., sp. nov., and S.waltheri Vagalinski, gen. nov., sp. nov.Colchiobrachyiulus Lohmander, 1936, a former subgenus of Megaphyllum, is here elevated to a full genus, and the genus Grusiniulus Lohmander, 1936 is downgraded to a subgenus of the genus Cyphobrachyiulus Verhoeff, 1900, both stat. nov., with their previously described species, Colchiobrachyiulusdioscoriadis (Lignau, 1915) and Cyphobrachyiulusredikorzevi (Lohmander, 1936), respectively, listed as comb. nov. Omobrachyiulusbrachyurus (Attems, 1899) is formally established as a junior subjective synonym of O.caucasicus (Karsch, 1881), syn. nov., and Omobrachyiulusimplicitusritsensis (Golovatch, 1981) is formally synonymised with the typical Omobrachyiulusimplicitus (Lohmander, 1936), syn. nov. Omobrachyiulussevangensis (Lohmander, 1932), originally described in the genus Megaphyllum, is here transferred to the former genus, comb. nov. The diagnoses and descriptions of some genera and subgenera are refined and complemented. A key is given to all genera and species of Brachyiulini that occur in the Caucasus, and their distributions are mapped. Several species are recorded as new to the faunas of Armenia, Azerbaijan, Georgia, or Russia. The distribution patterns of the Caucasian Brachyiulini and their biogeographic implications are discussed.

Rhodopotyphlus mitovi gen. et sp. nov., a new endogean millipede from the Western Rhodope Mountains, Bulgaria (Diplopoda: Julida: Julidae).

A new genus and species, Rhodopotyphlus mitovi gen. et sp. nov., is described from the Western Rhodopi Mts., south-central Bulgaria. According to its morphology combined with previously published molecular data, the new species is supposed to represent a basal lineage in the julid tribe Typhloiulini. Some assumptions concerning the early evolutionary stages of the development of the mesomerean important part of the male copulatory apparatus of many julid millipedesare made on the basis of the outstanding gonopod conformation in the newly described taxon. Rhodopotyphlus mitovi gen. et sp. nov. is likely a narrow local endemic showing preference for specific microhabitat conditions.

Revision of the millipede tribe Brachyiulini Verhoeff, 1909 (Diplopoda: Julida: Julidae), with descriptions of new taxa.

A comprehensive taxonomic revision of the circum-Mediterranean julid tribe Brachyiulini is undertaken. A total of 13 genera and 103 species are assigned to the tribe. One new genus, one new subgenus, and 10 new species are described from Greece, Turkey or Iran: Byzantorhopalum (Byzantorhopalum) elephantum sp. n., B. (Ioniulus) clavamum subgen. n., sp. n., Cyphobrachyiulus (Cyphobrachyiulus) spartanus sp. n., C. (Diaxylus) graecus sp. n., Graecoiulus skopelius gen. n., sp. n., Italoiulus poseidoni sp. n., Omobrachyiulus strasseri sp. n., Pontobrachyiulus falciferus sp. n., P. lohmanderi sp. n., Syriobrachyiulus golovatchi sp. n. 13 species are redescribed in detail, while the descriptions of another 36 species are complemented. The genera Anaulaciulus Pocock, 1895 and Mammamia Akkari et al., 2011 are formally excluded from the Brachyiulini. The following former subgenera of the genus MegaphyllumVerhoeff, 1894 are elevated to full genera: Byzantorhopalum Verhoeff, 1930, Cyphobrachyiulus Verhoeff, 1900, Iraniulus Attems, 1951, Italoiulus Attems, 1940, Omobrachyiulus Lohmander, 1936, Pontobrachyiulus Lohmander, 1936, Syriobrachyiulus Verhoeff, 1930. The genera Acropoditius Strasser, 1980 and Rhamphidoiulus Attems, 1905 are downgraded to subgenera of the genus Cyphobrachyiulus. The subgenera Phauloiulus Attems, 1940 and Persebrachyiulus Golovatch, 1983 are new subjective synonyms of Byzantorhopalum Verhoeff, 1930 and Syriobrachyiulus Verhoeff, 1930, respectively. The monophyly of the Brachyiulini in its current taxonomic composition is proven by morphological data, and supported by molecular and spermatological data in addition. The biogeographic affinities of the different tribal subgroups are briefly discussed.

Two new species of the millipede genus Metonomastus Attems, 1937 from the Balkan Peninsula (Diplopoda, Polydesmida, Paradoxosomatidae).

In addition to the eleven previously known species of the Mediterranean genus Metonomastus, two more species are described: M.petrovi sp. n., from the Rhodopi Mts. and Bunardzhik Hill in Bulgaria, and M.radjai sp. n., from the island of Mljet in Croatia. The relationships between the congeners and their distributions are briefly discussed. All 13 species of the genus are keyed.

Methyl N-methylanthranilate: major compound in the defensive secretion of Typhloiulus orpheus (Diplopoda, Julida).

The defensive secretion of the julid diplopod Typhloiulus orpheus contains methyl N-methylanthranilate (MNMA), an ester of N-methylanthranilic acid that comprises more than 99% of secretion of this species. MNMA is accompanied by small amounts of methyl anthranilate and two benzoquinones (2-methyl-1,4-benzoquinone and 2-ethyl-1,4-benzoquinone, respectively). MNMA is a known intermediate in the biosynthesis of both benzoquinones (as present in defensive secretions of juliformians) and glomerin-like quinazolines (chemical defense in Glomerida). The compound may have evolved independently in the pathway to glomeridan chemistry, or may even represent a pivotal branching point in the pathway to different chemical classes of diplopod defensive chemistry.

New records of millipedes and centipedes from Bulgaria, with an annotated checklist of the Bulgarian myriapods.

The present paper records new and poorly known myriapods (Diplopoda, Chilopoda) collected in Bulgaria in the last 10 years. Four new species are reported as new to the Bulgarian fauna: Lithobius melanops Newport, 1845 (Chilopoda: Lithobiomorpha: Lithodiidae), Polydesmus collaris C.L. Koch, 1847, Polydesmus inconstans Latzel, 1884 and Polydesmus schaessburgensis Verhoeff, 1898 (Diplopoda: Polydesmida: Polydesmidae). Scanning electron micrographs of the gonopods and male leg-pairs 5 and 6 are provided for the first time for Bulgarosoma superficiei Strasser, 1975, a species hitherto known only from its original description. This article presents also an updated list of Myriapoda in Bulgaria. The list includes a total of 251 (sub-)species of the classes Diplopoda (127), Chilopoda (104), Pauropoda (18) and Symphyla (2).

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.

"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.

A review of the millipede genus Typhloiulus Latzel, 1884 (Diplopoda: Julida: Julidae), with a description of three new species from Bulgaria and Greece.

The mostly cavernicolous and endogean millipede genus Typhloiulus Latzel, 1884 (Julida: Julidae) is hitherto known to comprise 33-36 species distributed in the Balkan and Apennine peninsulas, as well as the adjacent parts of the Alps and the Carpathian Arch. Here we describe three new species, T. bulgaricus sp. n., T. orpheus sp. n. and T. rhodopinus sp. n., collected from caves and other subterranean habitats in Bulgaria and Greece. Furthermore, the taxonomic descriptions of some poorly known species are emended and/or new distribution data given. T. staregai Strasser, 1973, is synonymized with T. strictus (Latzel, 1882). A neotype of T. kotelensis Jawlowski, 1938, of which the original type material is thought to be lost, is designated, and a checklist of Typhloiulus species is provided.

Millipedes of Cyprus (Myriapoda: Diplopoda).

The paper presents an annotated catalogue of the millipedes (Diplopoda) of Cyprus, based on literature scrutiny and on hitherto unpublished material. A total of 21 species belonging to 14 genera, 9 families and 7 orders are recorded from the island. Three species are regarded as new to science, but are not formally described, and the status of another three is yet to be clarified. Pachyiulus cyprius Brölemann, 1896 and Strongylosoma (Tetrarthrosoma) cyprium Verhoeff, 1902 are established as junior subjective synonyms of Amblyiulus barroisi (Porat, 1893) and T. syriacum (Humbert & DeSaussure, 1869), respectively, both syn. nov. The present-day diversity of the Cypriotic millipede fauna is analysed in the light of past geological events. It is presumed that the fauna originated from the late Miocene, when Cyprus was formed as a result of uplift of the sea floor. During the Messinian salinity crisis, some 6-5.3 mya, land bridges existed between the island and the neighbouring Taurus Mts. (Turkey) and the region of Latakia (Syria), which are supposed to have served as migratory pathways for various millipede taxa.

Redescription of the subgenus Parancistrum Verhoeff, 1943, an eastern Mediterranean lineage of the millipede genus Megaphyllum Verhoeff, 1894 (Diplopoda: Julida: Julidae: Brachyiulini).

The subgenus Parancistrum Verhoeff, 1943 is redefined on the basis of the examination of type and non-type material. The subgenus is currently settled to comprise 6 species, of which Megaphyllum arcuatum sp. n. is described as new, while M. adanense (Verhoeff, 1943) syn. n. and M. palaestinum (Jawlowski, 1931) syn. n. are established as junior subjective synonyms of M. curvifolii (Verhoeff, 1898) and M. genezarethanum (Verhoeff, 1923) comb. nov., respectively. A neotype is designed for Chromatoiulus tenenbaumi Jawlowski, 1931 under ICZN Article 75.1. The distribution pattern of the studied group is discussed, and some preliminary phylogenetic ideas about Megaphyllum and the tribe Brachyiulini are presented.

Redefinition of the millipede subgenus Megaphyllum sensu stricto Verhoeff, 1894 and neotype designation for Megaphyllum austriacum (Latzel, 1884) (Myriapoda: Diplopoda: Julida: Julidae).

Megaphyllum sensu stricto, i.e. the nominotypical subgenus of the very complicated genus Megaphyllum Verhoeff, 1894, is redefined on the basis of examination of type and non-type material and literature data. Four species-groups including twenty species in all are listed, and the identity of M. austriacum (C. L. Koch, 1838) is clarified with a proposal of neotype designation under ICZN Article 75.6. in order to stabilize the current usage of the name. Prevailing usage of M. silvaticum (Verhoeff, 1898) syn. nov. (nomen protectum) over the senior synonym M. nigrescens (Latzel, 1884) (nomen oblitum) is maintained under ICZN Article 23.9. M. banaticum (Verhoeff, 1899) is syn. nov. of M. erythronotum (Latzel, 1884) comb. nov., M. bosniense cotinophilum (Loksa, 1962) syn. nov. of M. bosniense bosniense (Verhoeff, 1897) and M. transsylvanicum transdanubicum (Loksa, 1962) syn. nov. of M. transsylvanicum transsylvanicum (Verhoeff, 1897). M. unilineatum (C. L. Koch, 1838) is new to the fauna of Turkey.