We present a revision of the genus Troglophilus in the western part of the Balkan Peninsula based on morphological and genetic analysis, and provide distribution data for the species. The results of these analyses reduced the number of the known taxa within the genus Troglophilus in this area to five valid species: Troglophilus cavicola (Kollar, 1833), Troglophilus neglectus Krauss, 1879, Troglophilus ovuliformis Karny, 1907, Troglophilus brevicauda Chopard, 1934, and Troglophilus lazaropolensis Karaman, 1958. A new species Troglophilus zorae sp. nov. is described and Troglophilus neglectus serbicus Maran, 1958, Troglophilus neglectus vlasinensis Maran, 1958, Troglophilus bukoviki Karaman, 1968 and Troglophilus pretneri Us, 1970 are synonymized. The distribution pattern of the western Balkan species supports a general east-west migration route. We also determined the centres of origin and spreading directions of particular species according to the distribution of the taxa and the structure of the haplotypes.
Large prosomal scent glands constitute a major synapomorphic character of the arachnid order Opiliones. These glands produce a variety of chemicals very specific to opilionid taxa of different taxonomic levels, and thus represent a model system to investigate the evolutionary traits in exocrine secretion chemistry across a phylogenetically old group of animals. The chemically best-studied opilionid group is certainly Laniatores, and currently available chemical data allow first hypotheses linking the phylogeny of this group to the evolution of major chemical classes of secretion chemistry. Such hypotheses are essential to decide upon a best-fitting explanation of the distribution of scent-gland secretion compounds across extant laniatorean taxa, and hence represent a key toward a well-founded opilionid chemosystematics.
While considerable knowledge on the chemistry of the scent gland secretions from the opilionid suborders Laniatores and Cyphophthalmi has been compiled, it is the Palpatores (Eupnoi and Dyspnoi) where chemical data are scarce. In particular, the Dyspnoi have remained nearly unstudied, mainly due to their reported general reluctance to release secretions as well as to the phenomenon of production of insoluble—and inaccessible—solid secretion. We here show that at least certain nemastomatid Dyspnoi, namely all three species of genus Carinostoma, indeed produce a volatile secretion, comprising octan-3-one, 6-methyl-5-hepten-2-one and acetophenone in species-specific combinations. In all Carinostoma spp., these volatiles are embedded in a semi-volatile, naphthoquinone matrix (mainly 1,4-naphthoquinone and 6-methyl-1,4-naphthoquinone). In detail, acetophenone and traces of naphthoquinones characterize the secretions of Carinostoma carinatum. A mixture of octan-3-one, 6-methyl-5-hepten-2-one and large amounts of naphthoquinones were found in C. elegans, and 6-methyl-5-hepten-2-one together with small amounts of naphthoquinones in the secretions of C. ornatum. So far, exclusively naphthoquinones had been reported from a single dyspnoan hitherto studied, Paranemastoma quadripunctatum.
Aim To investigate the phylogeny of the genus Cyphophthalmus in the Balkan Peninsula and to test the current recognition of ‘phyletic lines’ and phylogenetic groups proposed in previous studies in order to elucidate the biogeographical history of the region.
Location Europe, Balkan Peninsula, Adria microplate.
Methods Two mitochondrial (cytochrome c oxidase subunit I and 16S rRNA) and two nuclear (28S rRNA and 18S rRNA) markers were used to infer the phylogenetic history of the group. Molecular dating with relaxed molecular clocks was used to elucidate the relative time of diversification within the genus Cyphophthalmus and its constituent lineages.
Results Our analyses confirm the monophyly of the genus Cyphophthalmus, and that of the Aegean and gjorgjevici lineages, whereas the ‘Dinaric lineage’ appears paraphyletic.
Main conclusions We show that the genus Cyphophthalmus is an old endemic from the Balkan biogeographical region, which gave origin to at least three main lineages. Those lineages have diversified within overlapping ranges. According to our molecular dating, they have also diversified within the same timeframe. The Dinaric Alps, although presenting a large number of species, cannot be inferred as the centre of origin of the group. Instead, the biogeographical evolution of the genus could be related to the palaeogeographic history of the Adria microplate.
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