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.
Diplura is a group of entognathous hexapods, often considered a sister group to insects. They play an important role in recycling organic matter in soil and subterranean terrestrial ecosystems. The Campodeidae is the most diverse family, divided into four subfamilies. The subfamily Plusiocampinae has a subterranean life-style with many species distributed in the Euro-Mediterranean area. The incertae sedis tachycampoids (“lignée Tachycampoïde”) is a group within the family Campodeidae that share with the Plusiocampinae a strong preference for subterranean habitats and several morphological characters, such as slender body shape, elongated appendages, considerable increment in the number of antennomeres and cercal articles, and complexity of sensorial structures. The present monograph provides a taxonomic revision of the subfamily Plusiocampinae and the genera belonging to the tachycampoid lineage from Europe and the Mediterranean region. It comprises detailed morphological descriptions and illustrations together with data on the habitats and distributions of 87 species, 10 subspecies and 11 affinis forms. Seven new species are described among those, namely: Plusiocampa (Plusiocampa) apollo Sendra, Giachino & Vailati sp. nov., P. (P.) chiosensis Sendra & Gasparo sp. nov., P. (P.) dublanskii Sendra & Turbanov sp. nov., P. (P.) hoffmanni Sendra & Paragamian sp. nov., P. (P.) rhea Sendra sp. nov., P. (P.) ternovensis Sendra & Borko sp. nov. and P. (Venetocampa) ferrani Sendra & Delić sp. nov.
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.Electronic supplementary materialThe online version of this article (doi:10.1007/s10886-016-0680-4) contains supplementary material, which is available to authorized users.
The Caucasian leucogeorgiinine genera Archileucogeorgia Lohmander, 1936 and Leucogeorgia Verhoeff, 1930 are revised, with Leucogeorgia being considered as a senior subjective synonym of Archileucogeorgia, syn. nov. The following new combinations are thus warranted: Leucogeorgia abchasica (Lohmander, 1936) and L. satunini (Lohmander, 1936), both comb. nov. ex Archileucogeorgia. All four previously described species, viz., L. longipes Verhoeff, 1930, L. abchasica, L. satunini and L. rediviva Golovatch, 1983, are redescribed based on new material, partly also on the type series, with a lectotype being designated for L. longipes. In addition, eleven new species of Leucogeorgia are described, both with normal (L. borealis sp. nov., L. gioi sp. nov., L. golovatchi sp. nov., L. lobata sp. nov., L. oculata sp. nov. and L. prometheus sp. nov.) and modified mouthparts (L. caudata sp. nov., L. mystax sp. nov., L. profunda sp. nov., L. redivivoides sp. nov. and L. turbanovi sp. nov., all clearly troglobionts). Additionally, a new monotypic genus, Martvilia gen. nov., is erected, with M. parva gen. et sp. nov. as the type species, another presumed troglobiont. An identification key to both genera and all 16 species of Caucasian Leucogeorgiini is presented. Several other members of this tribe are also re-examined, including a syntype male of Telsonius nycteridonis Strasser, 1976, from Greece, herewith designated as the lectotype. Troglomorphisms and mouthpart modifications, as well as the distributions and relationships within both Leucogeorgia and Leucogeorgiini, and a Leucogeorgiini species richness estimate for the western Caucasus are discussed.
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.Electronic supplementary materialThe online version of this article (doi:10.1007/s10886-017-0832-1) contains supplementary material, which is available to authorized users.
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