A new dwarf schendylid centipede (Chilopoda: Geophilomorpha: Schendylidae) with a low number of legs from Serbia, Balkan Peninsula.

Schendyla antici sp. nov., a dwarf geophilomorph from the Mt. Medvednik (Western Serbia, Balkan Peninsula), is described and illustrated based on the specimens extracted from the soil samples. A detailed comparison with all species within the genus is provided. The new species has the lowest number of leg-bearing segments within the genus Schendyla Bergse & Meinert, 1866, and one of the lowest in the order Geophilomorpha in general.

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.

Life after the mother's hug: Late post-embryonic development of Cryptops parisi (Chilopoda: Scolopendromorpha: Cryptopidae).

Here we describe in detail the late post-embryonic development of the common European scolopendromorph centipede Cryptops parisi. Canonical variate analyses of two groups of external morphological characters, viz., cephalic capsule characters (head length, length of the anterior and posterior paramedian cephalic sutures) and coxopleuron surface characters (number of pores in the coxal pore-field, number of setae on the posterior coxopleuron edge, their number on the coxal pore-field, and their number posterior to the coxal pore-field) were conducted on a large sample of specimens collected from two localities in Serbia. Ten free-living stages are recognized: three pre-adult stages (adolescens I, II, and III) and seven adult stages (one maturus junior stage, four maturus, and two maturus senior stages). The fourth late post-embryonic stage is the first mature stage in both sexes. Sexual dimorphism in the aforementioned characters was not observed. Morphological variation of coxopleuron characters was more informative for the discrimination of developmental stages in Cryptops than the morphological variation of cephalic capsule characters.

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.

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.

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.

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 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.

Review of the family Anthogonidae (Diplopoda, Chordeumatida), with descriptions of three new species from the Balkan Peninsula.

Three new species of the family Anthogonidae are described from caves in Croatia and Montenegro, respectively: Egonpretneria vudutschajldi Antic & Drazina sp. n., Haasia jalzici Antic & Drazina sp. n., and Macrochaetosoma bertiscea Antic & Makarov sp. n. All three subfamilies within Anthogonidae are diagnosed, with brief discussion of relationships between genera and notes on their biogeography. A map of the global distribution of Anthogonidae is presented, as well as maps of the distribution of each species of Balkan anthogonids, including numerous new locality data. The first SEM images are provided for some representatives of this family. A key is given to all seven anthogonid genera. The subfamily Haasiinae Hoffman, 1980 is a new synonym of Anthogonidae, syn. n., while Macrochaetosoma bifurcatum Curcic & Makarov, 2001 becomes a new synonym for M. troglomontanum Absolon & Lang, 1933, syn. n..

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.

Micro- and nanostructures of iridescent wing scales in purple emperor butterflies (Lepidoptera: Apatura ilia and A. iris).

Apatura ilia (Denis and Schiffermüller, 1775) and A. iris (Linnaeus, 1758) are fascinating butterflies found in the Palaearctic ecozone (excepting the north of Africa). The wings of these insects are covered with a great number of two types of scales positioned like roof tiles. Type I scales are on the surface, while type II scales are situated below them. The structural color of the type I scales is recognized only on the dorsal side of both the fore and hind wings of the males of the aforementioned species. Both types of scales are responsible for pigment color of the wings, but iridescence is observed only in the type I scales. The brilliant structural color is due to a multilayer structure. The features of the scales, their dimensions and fine structure were obtained using scanning electron microscopy. Cross sections of the scales were then analyzed by transmission electron microscopy. The scales of the "normal" and clytie forms of A. ilia have a different nanostructure, but are of the same type. A similar type of structure, but with a different morphology, was also noticed in A. iris. The scales of the analyzed species resemble the scales of tropical Morpho butterflies.

Composition of the defensive secretion in three species of European millipedes.

Three European julid species, Cylindroiulus boleti, Leptoiulus trilineatus, and Megaphyllum bosniense, secrete mixtures of up to 12 different quinones. The major components in these species are 2-methoxy-3-methyl-1,4-benzoquinone and 2-methyl-1,4-benzoquinone. 2-Methoxy-5-methylhydroquinone is detected for the first time in the Class Diplopoda. 2-Hydroxy-3-methyl-1,4-benzoquinone, 2,3-dimethoxyhydroquinone, 2-methyl-3,4-methylendioxyphenol, and 2,3-dimethoxy-5-methylhydroquinone are registered for the first time in representatives of the family Julidae. The similar chemical composition of defense secretions in all analyzed European julids and Pacific spirobolids supports the idea of the chemical consistency of defensive compounds in juliform millipedes.

Defensive secretions in Callipodella fasciata (Latzel, 1882; Diplopoda, Callipodida, Schizopetalidae).

The small millipede Callipodella fasciata secretes an earthy smell when disturbed. This secretion was obtained by CH(2) Cl(2) extraction from specimens of both sexes and was identified by GC/MS analyses to be composed of p-cresol (96.5%), phenol (3.5%), and p-ethylphenol (traces). This is the first identification of these compounds in an epigean European callipodidan species and the first report of intergeneric differences in the chemical composition of defensive secretions in callipodidans. These compounds have repellent, antimicrobial, and antifungal properties.

Defensive secretions in three species of polydesmids (Diplopoda, Polydesmida, Polydesmidae).

Nine compounds were detected in three different millipede species: Polydesmus complanatus (L.), Brachydesmus (Stylobrachydesmus) avalae Curcic & Makarov, and Brachydesmus (Stylobrachydesmus) dadayi Verhoeff. Benzaldehyde, benzyl alcohol, benzoylnitrile, benzyl methyl ketone, benzoic acid, benzyl ethyl ketone, mandelonitrile, and mandelonitrile benzoate were identified by GC-FID and GC-MS analyses. Hydrogen cyanide was detected qualitatively by the picric acid test. Benzyl ethyl ketone, benzyl methyl ketone, and benzyl alcohol were detected for the first time in polydesmidan millipedes. Benzoylnitrile was the major component in all three hexane extracts. These compounds are suspected to be active in the defensive secretions of these millipede species.

Identification of secretory compounds from the European callipodidan species Apfelbeckia insculpta.

Defensive secretions of the callipodidan species Apfelbeckia insculpta contain a p-cresol as the main component and phenol in traces. This is the first identification of these compounds in a European callipodidan species. The repugnatory glands of A. insculpta are of the spirobolid type and consist of a spherical reservoir, a simple duct, and a valvular cuticular infolding that opens onto the lateral surface of the millipede via a pore.