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.

Dazbogosoma mokoshae, a new cavernicolous species of the previously monospecific genus Dazbogosoma Makarov amp; uri, 2012, from Serbia (Diplopoda: Chordeumatida: Anthroleucosomatidae).

A new troglobiotic species, Dazbogosoma mokoshae sp. nov., is described from a cave in the eastern Serbia. This is the second species from the previously monospecific genus Dazbogosoma Makarov uri, 2012. An updated diagnosis of the genus, as well as a diagnosis, detailed description, illustrations and in situ photographs of a new species, based on both males and females, are presented. The distribution and relationships within the Bulgarosoma complex comprising D. mokoshae sp. nov. are briefly discussed.

Morphological integration of the head capsule in the millipede Megaphyllum unilineatum (C. L. Koch, 1838) (Diplopoda: Julida): can different modules be recognized?

Covariation of multiple morphological traits and modularity have been widely studied in the field of evolutionary developmental biology. Subunits of a morphological structure can evolve separately from each other in a modular fashion. The aims of our study therefore were: i) to test the hypothesis of modularity in the dorsal part of the head capsule and the gnathochilarium separately during late postembryogenesis in the julidan millipede Megaphyllum unilineatum (C. L. Koch, 1838) using geometric morphometrics; and ii) to investigate the influence of allometry on overall morphological integration in the dorsal part of the head capsule and the gnathochilarium in the mentioned species. Individuals from different ontogenetic stadia (stadium VI - stadium XI) were included in the analyses. Significant influence of fluctuating asymmetry on the dorsal part of the head capsule shape was detected by Procrustes ANOVA. Regressions were significant for the symmetric component of both analysed morphological traits, while non-significant regression was detected for the asymmetric component of the head capsule's dorsal part. Hypotheses of modularity for the dorsal part of the head capsule and the gnathochilarium are rejected because our results indicate that a small proportion of alternate partitions has higher covariation between subsets of structure than between the hypothesized modules. Contrary to our expectations, results of the present study show that allometry does not increase the level of morphological integration in the dorsal part of the head capsule and the gnathochilarium in M. unilineatum. Based on the obtained results, we conclude that the dorsal part of the head capsule and the gnathochilarium are not composed of independent modules and that in the case of the capsule's dorsal part, developmental processes affect morphological integration in different ways at different levels of shape variation.

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.

Sexual size and shape dimorphism in Brachydesmus troglobius Daday, 1889 (Diplopoda, Polydesmida).

Until now, morphological trait variation has been investigated in several millipede species using geometric morphometrics. The present study is the first attempt to explore sexual shape and size dimorphism (SShD and SSD) of morphological structures in Polydesmida. We here analyse antennal, head, and leg SShD and SSD in Brachydesmus troglobius Daday, 1889. Our results show that SSD exists in all of the analysed structures, while SShD is present only in the legs. In comparison with females, males possess longer and wider legs, as well as longer antennae and a shorter head. Contrary to previous findings in some Julida, in B. troglobiusSSD of the antennae and legs varies more than SShD in these morphological structures.

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.

Phenol-based millipede defence: antimicrobial activity of secretions from the Balkan endemic millipede Apfelbeckia insculpta (L. Koch, 1867) (Diplopoda: Callipodida).

Millipedes use an array of chemical compounds to defend themselves from predator attack. These chemical substances can have additional roles, i.e. defence against various pathogens. We evaluated the efficacy of the defensive secretion of Apfelbeckia insculpta (L. Koch, 1867) against bacteria, yeasts, and filamentous fungi. The tested secretion consisted of two compounds, p-cresol and phenol, and showed antibacterial, antibiofilm, and antifungal potential against all selected microorganisms. The most sensitive bacterium in our study was Pseudomonas aeruginosa, while the tested defensive secretion manifested the lowest activity against Escherichia coli. The defensive secretion of A. insculpta also showed an ability, albeit mild, to suppress biofilm formation by P. aeruginosa. Among the tested yeasts, Candida albicans and C. krusei were the most susceptible and most resistant species, respectively. Finally, the concentration of extracts obtained from the tested defensive secretion needed to achieve an antifungal effect was lowest in the case of Cladosporium cladosporioides. Fusarium verticillioides and Penicillium rubens were the micromycetes most resistant to the tested secretion. Our results indicate that antibacterial activity of the defensive secretion of A. insculpta is similar to or slightly weaker than that of streptomycin, while comparison with antimycotics showed that the tested millipede secretion has stronger activity than fluconazole, but weaker activity than nystatin and ketoconazole. The present study corroborates previous findings indicating that the defensive secretions of millipedes can have different roles apart from antipredator protection and are effective against pathogenic microorganisms.

Multifaceted activity of millipede secretions: Antioxidant, antineurodegenerative, and anti-Fusarium effects of the defensive secretions of Pachyiulus hungaricus (Karsch, 1881) and Megaphyllum unilineatum (C. L. Koch, 1838) (Diplopoda: Julida).

Members of the millipede order Julida rely on dominantly quinonic defensive secretions with several minor, non-quinonic components. The free radical-scavenging activities of ethanol, methanol, hexane, and dichloromethane extracts of defensive secretions emitted by Pachyiulus hungaricus (Karsch, 1881) and Megaphyllum unilineatum (C. L. Koch, 1838) were investigated using the ABTS, DPPH, and total reducing power (TRP) tests. The obtained extracts were also tested for inhibition of acetylcholinesterase and tyrosinase activity. Finally, the antifungal potential of both julid extracts was evaluated against seven Fusarium species. Secretions of both species showed activity against free radicals, acetylcholinesterase, tyrosinase, and all of the selected fungal species. The secretions of P. hungaricus exhibited a more potent antioxidative effect than did those of M. unilineatum, while there were no significant differences of antiacetylcholinesterase activity between the tested extracts. Only the hexane extract of M. unilineatum showed an effect on tyrosinase activity stronger than that of P. hungaricus. Fusarium sporotrichioides, F. graminearum, and F. verticillioides were the fungi most resistant to secretions of both julids. The Fusarium species most susceptible to the secretion of P. hungaricus was F. avenaceum, while the concentrations of M. unilienatum extracts needed to inhibit and completely suppress fungal growth were lowest in the case of their action on F. lateritium. Our data support previous findings that julid defensive secretions possess an antimicrobial potential and reveal their antioxidative and antineurodegenrative properties. Bearing in mind the chemical complexity of the tested defensive secretions, we presume that they can also exhibit other biological activities.

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.

'Does my Diet Affect my Perfume?' Identification and Quantification of Cuticular Compounds in Five Drosophila melanogaster Strains Maintained over 300 Generations on Different Diets.

Cuticular hydrocarbons (CHCs) in Drosophila melanogaster represent the basis of chemical communication being involved in many important biological functions. The aim of this study was to characterize chemical composition and variation of cuticular profiles in five D. melanogaster strains. These strains were reared for approximately 300 generations on five diets: standard cornmeal medium and substrates prepared with apple, banana, tomato, and carrot. Differences in quantity and/or quality in CHCs were assumed as a result of activation of different metabolic pathways involved in food digestion and adaptations to the particular diet type. In total, independently of sex and strain, 66 chemical compounds were identified. In females of all strains, 60 compounds were identified, while, in males, 47 compounds were extracted. Certain new chemical compounds for D. melanogaster were found. MANOVA confirmed that CHC amounts significantly depend on sex and substrates, as well as on their interactions. Discriminant analysis revealed that flies belonging to 'apple' and 'carrot' strains exhibited the most noticeable differences in CHC repertoires. A non-hydrocarbon pheromone, cis-vaccenyl acetate (cVA) also contributed to the variation in the pheromone bouquet among the strains. Variability detected in CHCs and cVA may be used in the explanation of differences in mating behaviour previously determined in analyzed fly strains.

Molecular Diversity of Compounds from Pygidial Gland Secretions of Cave-Dwelling Ground Beetles: The First Evidence.

Three adult cave-dwelling ground beetle species were induced to discharge secretions of their pygidial glands into vials. Dichloromethane extraction was used to obtain the secretions. In total, 42 compounds were identified by GC/MS analysis. Pheggomisetes ninae contained 32 glandular compounds, Laemostenus (Pristonychus) punctatus 13, whereas Duvalius (Paraduvalius) milutini had nine compounds. Caproic, oleic, palmitic, and stearic acids were present in the samples of all analyzed species. Undecane was predominant in the extract of L. punctatus. Palmitic acid was the major component in the secretion of D. milutini. Finally, the most abundant compounds in P. ninae secretion were heptacosene and nonacosadienes. Herein, we present the first data on the identification of pygidial gland secretion components in both troglophilous and troglobite cave-dwelling ground beetles. Some compounds are reported for the first time in the secretions of ground beetles and other higher or lower taxa. The adaptation to underground life has not led to a reduction or changes in the chemical defense mechanism in the analyzed troglophilous and troglobitic Platyninae and Trechinae taxa.

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.