Mladen Kučinić

Pleistocene divergence of Dinaric Drusus endemics (Trichoptera, Limnephilidae) in multiple microrefugia within the Balkan Peninsula

The Balkan Peninsula is one of three major European refugial areas. It has high biodiversity and endemism, but data on the age and origin of its fauna, especially endemics, are limited. Mitochondrial sequence data (COI and 16S genes) were used to study the population structure and phylogeography of the caddisfly Drusus croaticus and the phylogeny and divergence of seven other Drusus species, mostly range‐restricted endemics of the Dinaric region of the Balkan Peninsula. The divergence of D. croaticus populations in Croatia and allopatric Drusus species in Bosnia dated to the Pleistocene, showing the importance of this time period for the origin and diversification of Balkan endemic taxa. The divergence of more distantly related species dated to the Late Miocene/Early Pliocene. Population genetic and phylogeographic analysis of 115 individuals from 11 populations of D. croaticus revealed a high level of genetic differentiation and absence of gene flow between populations separated by more than 10 km. The existence of allopatrically fragmented lineages in D. croaticus and the endemic Bosnian species is most likely the result of long‐term isolation in multiple microrefugia, probably due to the specific habitat requirements and life‐history traits of Drusinae coupled with the topographic complexity and historical changes in geomorphology of the region. Overall, these findings shed light on the processes generating the high genetic complexity of this refugial region that parallels the ‘refugia within refugia’ pattern widely reported from the Iberian refugium.

Microscale vicariance and diversification of Western Balkan caddisflies linked to karstification

Abstract: The karst areas in the Dinaric region of the Western Balkan Peninsula are a hotspot of freshwater biodiversity. Many investigators have examined diversification of the subterranean freshwater fauna in these karst systems. However, diversification of surface-water fauna remains largely unexplored. We assessed local and regional diversification of surface-water species in karst systems and asked whether patterns of population differentiation could be explained by dispersal—diversification processes or allopatric diversification following karstrelated microscale vicariance. We analyzed mitochondrial cytochrome c oxidase subunit I (mtCOI) sequence data of 4 caddisfly species (genus Drusus) in a phylogeographic framework to assess local and regional population genetic structure and Pliocene/Pleistocene history. We used BEAST software to assess the timing of intraspecific diversification of the target species. We compared climate envelopes of the study species and projected climatically suitable areas during the last glacial maximum (LGM) to assess differences in the species climatic niches and infer potential LGM refugia. The haplotype distribution of the 4 species (324 individuals from 32 populations) was characterized by strong genetic differentiation with few haplotypes shared among populations (16%) and deep divergence among populations of the 3 endemic species, even at local scales. Divergence among local populations of endemics often exceeded divergence among regional and continental clades of the widespread D. discolor. Major divergences among regional populations dated to 2.0 to 0.5 Mya. Species distribution model projections and genetic structure suggest that the endemic species persisted in situ and diversified locally throughout multiple Pleistocene climate cycles. The pattern for D. discolor was different and consistent with multiple invasions into the region. Patterns of population genetic structure and diversification were similar for the 3 regional endemic Drusus species and consistent with microscale vicariance after the onset of intensified karstification in the Dinaric region. Karstification may induce microscale vicariance of running surface-water habitats and probably promotes allopatric fragmentation of stream insects at small spatial scales.

The caddisfly fauna (Insecta, Trichoptera) of the rivers of the Black Sea basin in Kosovo with distributional data for some rare species

Abstract Adult caddisflies were collected from 12 stations in the Black Sea basin in Kosovo using UV light traps. Sixty-five of the seventy-six species reported in this paper are first records for the Kosovo caddisfly fauna. The unexpected discovery of several species during this investigation: Agapetus delicatulus McLachlan, 1884, Psychomyia klapaleki Malicky, 1995, Tinodes janssensi Jacquemart, 1957, Hydropsyche emarginata Navas, 1923, Drusus botosaneanui Kumanski, 1968, Potamophylax rotundipennis (Brauer, 1857), Potamophylax schmidi Marinković-Gospodnetić, 1970, Ceraclea albimacula (Rambur, 1842), Helicopsyche bacescui Orghidan & Botosaneanu, 1953, Adicella filicornis (Pictet, 1834), Beraea maurus (Curtis, 1834) and Beraeamyia hrabei Mayer, 1937 illustrates that collections from poorly investigated areas in Europe will almost certainly revise the existing knowledge on the distribution of these and other species.

A cave leech (Hirudinea, Erpobdellidae) from Croatia with unique morphological features

Croatobranchus mestrovi is a troglobitic leech from deep shaft‐like caves in the Velebit Mountain, Dinaric karst, Croatia, living in cold (4–6 °C) water. Its oral sucker extends to form four pairs of triangular tentacles, each with about five finger‐like papillae, but widening into a marginally crenulated disc when attached to a substrate. Pairs of stiff, finger‐like lateral projections, probably gills, occur along the body behind the clitellum. Somites are simple five‐annulate. Despite the unique head morphology and the presence of lateral outgrowths, the anatomy and 18S rRNA gene sequence of this species indicate that it is a member of the Erpobdellidae, closely related to Dina.

Description of a new species of Wormaldia from Sardinia and a new Drusus species from the Western Balkans (Trichoptera, Philopotamidae, Limnephilidae)

Abstract New species are described in the genera Wormaldia (Trichoptera, Philopotamidae) and Drusus (Trichoptera, Limnephilidae, Drusinae). Additionally, the larva of the new species Drusus crenophylax sp. n. is described, and a key provided to larval Drusus species of the bosnicus-group, in which the new species belongs. Observations on the threats to regional freshwater biodiversity and caddisfly endemism are discussed. The new species Wormaldia sarda sp. n. is an endemic of the Tyrrhenian island of Sardinia and differs most conspicuously from its congeners in the shape of segment X, which is trilobate in lateral view. The new species Drusus crenophylax sp. n. is a micro-endemic of the Western Balkans, and increases the endemism rate of Balkan Drusinae to 79% of 39 species. Compared to other Western Balkan Drusus, males of the new species are morphologically most similar to Drusus discophorus Radovanovic and Drusus vernonensis Malicky, but differ in the shape of superior and intermediate appendages. The females of Drusus crenophylax sp. n. are most similar to those of Drusus vernonensis, but differ distinctly in the outline of segment X. Larvae of Drusus crenophylax sp. n. exhibit toothless mandibles, indicating a scraping grazing-feeding ecology.

Carabid beetle diversity and mean individual biomass in beech forests of various ages.

Abstract Carabid beetle diversity and mean individual biomass (MIB) were analysed in three different successional stages of beech tree stands (60, 80 and 150 years old). Carabid beetles were captured using pitfall traps placed at nine sites (three per age class) in the Papuk Mountain of East Croatia during 2008. A cluster analysis identified three groupings that corresponded to the beech age classes. MIB values increased with stand age, ranging from 255 in 60-year-old stand to 537 in the oldest forests. The 80-year-old stand showed the highest species richness and diversity values. With respect to species composition, large species such as Carabus scheidleri and Carabus coriaceus were dominant only in the oldest forests. Furthermore, species that overwinter in the larval stage were more abundant in the oldest forests (45% of the total number of individuals from the 150-year-old stand) than in the younger ones (20% of individuals from 60-year-old, and 22% of individuals from 80-year-old stands). Our results showed that the analyses of species composition and life history traits are valuable for estimating the conservation values of older forests. Although the investigated sites form part of a continuous forested area and are only a couple of kilometres apart, MIB values detect significant differences associated with forest age and can be a useful tool in evaluating the degree to which a forest reflects a natural state.

The larva and life cycle of Annitella apfelbecki (Klapalek, 1899), including a redescription of Melampophylax nepos (McLachlan, 1880) (Trichoptera: Limnephilidae)

The hitherto unknown larva of Annitella apfelbecki (Klapálek, 1899) is described and discussed in the context of contemporary Limnephilidae keys. In addition, its life cycle is discussed, and zoogeographical and ecological notes are included. Finally, we provide some additional information on the morphologically very similar larva of Melampophylax nepos (McLachlan, 1880).

A hairy case: The evolution of filtering carnivorous Drusinae (Limnephilidae, Trichoptera)

The caddisfly subfamily Drusinae BANKS comprises roughly 100 species inhabiting mountain ranges in Europe, Asia Minor and the Caucasus. A 3-gene phylogeny of the subfamily previously identified three major clades that were corroborated by larval morphology and feeding ecologies: scraping grazers, omnivorous shredders and filtering carnivores. Larvae of filtering carnivores exhibit unique head capsule complexities, unknown from other caddisfly larvae. Here we assess the species-level relationships within filtering carnivores, hypothesizing that head capsule complexity is derived from simple shapes observed in the other feeding groups. We summarize the current systematics and taxonomy of the group, clarify the systematic position of Cryptothrix nebulicola, and present a larval key to filtering carnivorous Drusinae. We infer relationships of all known filtering carnivorous Drusinae and 34 additional Drusinae species using Bayesian species tree analysis and concatenated Bayesian phylogenetic analysis of 3805bp of sequence data from six gene regions (mtCOI5-P, mtCOI3-P, 16S mrDNA, CADH, WG, 28S nrDNA), morphological cladistics from 308 characters, and a total evidence analysis. All analyses support monophyly of the three feeding ecology groups but fail to fully resolve internal relationships. Within filtering carnivores, variation in head setation and frontoclypeus structure may be associated with progressive niche adaptation, with less complex species recovered at a basal position. We propose that diversification of complex setation and frontoclypeus shape represents a recent evolutionary development, hypothetically enforcing speciation and niche specificity within filtering carnivorous Drusinae.

The larva, ecology and distribution of Tinodes braueri McLachlan 1878 (Trichoptera: Psychomyiidae)

The larva of Tinodes braueri McLachlan 1878 is described for the first time and compared with the larvae of other species in the genus. In addition, notes on the zoogeography and ecology of the species are included.

Description of two new filtering carnivore Drusus species (Limnephilidae, Drusinae) from the Western Balkans

Abstract Two new species of the genus Drusus (Trichoptera, Limnephilidae, Drusinae) from the Western Balkans are described. Additionally, observations on the biodiversity and threats to the region’s endemic aquatic fauna are discussed. Drusus krpachi sp. n. is a micro-endemic of the Korab Mountains, Macedonia, and Drusus malickyi sp. n. is a micro-endemic of the Prokletije Mountains, Albania. Both new species are most similar to Drusus macedonicus but differ from the latter in the shape of segment IX, the shape of the tips of the intermediate appendages in lateral view, the shape of the inferior appendages, and the form and shape of the parameres. In addition, males of the European species of filtering carnivore Drusinae are diagnosed and illustrated, including Cryptothrix nebulicola McLachlan, Drusus chrysotus Rambur, Drusus discolor Rambur, Drusus macedonicus Schmid, Drusus meridionalis Kumanski, Drusus muelleri McLachlan, Drusus romanicus Murgoci and Botosaneanu, and Drusus siveci Malicky. These additions to the Western Balkan fauna demonstrate the significance of this region for European biodiversity and further highlight the importance of faunistic studies in Europe.