Search

The caddisfly Drusus dudor Oláh, 2017 (Limephilidae: Drusinae) was described from the Northwestern Italian Alps. We provide a detailed description of the larva, based on material from the Italian Province of Piemonte. Information on the morphology of the 5th larval instar is given, and the most important diagnostic features are illustrated. The larva is included in an updated key to larval Drusinae where D. dudor keys together with Drusus aprutiensis Moretti, 1981, D. camerinus Moretti, 1981, D. croaticus Marinkovic-Gospodnetic, 1971, D. mixtus (Pictet, 1834), and D. nigrescens Meyer-Duer, 1875. The species can be reliably separated by the morphology of the pronotum, the shape of the metanotal sclerites, and by morphological details of abdominal sternum I.

In a recent 3-gene phylogeny of the trichopteran subfamily Drusinae Banks 1916, molecular data clearly correlated with the morphology and feeding ecology of larvae. The largest of three main groups, the Drusinae grazer clade, exhibits an unusual larval feeding ecology for Limnephilidae, and is the most diverse group. In this paper we describe four previously unknown Drusinae larvae included in this clade: Drusus balcanicus Kumanski, 1973 (micro-endemic to Eastern Balkans), Drusus botosaneanui Kumanski, 1968 (Dinaric Western Balkans, Hellenic and Eastern Balkan, Asia Minor), Drusus serbicus Marinković-Gospodnetić, 1971 (micro-endemic to Dinaric Western Balkans), and Drusus tenellus (Klapálek, 1898) (Carpathians, Dinaric Eastern Balkans). Characteristically, the larvae of these species have toothless mandibles typical of the Drusinae grazer clade. Larvae and adults were unambiguously associated using a phylogenetic analysis based on two mitochondrial [mtCOI, mtLSU (=16S) rDNA] and two nuclear genes (nuWG, nuCAD). In addition, information on the morphology of the larvae is given and the diagnostic features necessary for identification are illustrated.

In the last few years we have described over 70 new incipient sibling limnephild species applying the discovered Trichoptera speciation traits of the paraproct and paramere for species recognition and delimitation. In this revision on Drusinae subfamily, comprising 177 species, we have applied these subtle, but rapid and stable speciation traits and described 49 new sibling species from the “well studied” European mountain ranges. Discussing the theoretical background we have elaborated and adapted a new character state ranking system of phenomics to revise the long-neglected taxonomy of the Drusinae subfamily and synonymised the Cryptothrix, Monocentra, Metanoea, Leptodrusus, Anomalopterygella, Hadimina genera with the Drusus genus. These old genera of artificial constructs were established exclusively by divergences of secondary sexual traits known already to have only species level ranking value. According to our new character ranking system in the Drusinae subfamily, beside the Drusus genus, only the Ecclisopteryx genus has been retained having robust generic level divegences of paraproct loss and ancestral duplication of spine organising centre on the paramere pattern. Speciation trait function of the peg-packed surface on the paraproct head in Drusus genus moved to the gonopod apices and integrated into variously shaped stimulatory organ in the Ecclisopteryx genus. In the Drusus genus the ancestral divergence of the single spine organising centre has integrated 11 species groups with remarkably stable paramere spine pattern. Based upon ancestral divergences in the paraproct architecture we have differenciated 28 species complexes inside the 11 species groups. The delineation of the 163 mostly incipient siblings species, inside the 28 species complexes with 44 new Drusus species, was based primarily on the divergences of speciation trait, that is in the stimulatory head shape of the apical arms on the dorsal branches of the paraproct. In the Ecclisopteryx genus with 14 species we have established two independent lineages both with a single species, as well as two species complexes with five new species applying the speciation trait of the genus, that is the shape divergence of the stimulatory organ on the dorsoapical surface of the gonopods. Based on the Darwinian natural selection, we do not understand how the discovered 70+49 new European incipient phylogenetic species of limnephilid caddisflies have been evolved in the isolated sky island habitats of high mountain ranges. This isolation induced speciation represents a challenge to the mechanistic reductionist concept of the natural selection. Our first trial to extract information from various disciplines to answer this question is presented in a brief theoretical discourse: (1) rethinking the status of natural selection towards postdarwinism; (2) teleology or teleonomy; (3) limits and potentials in understanding reality; (4) organisation of universe by integration; (5) what are and how the organising forces are powered to work in the emerging energy mechanisms; (6) divergence by integration; (7) divergence in isolation; (8) reproductive isolation by sexual selection; (9) shape divergence; (10) speciation traits; (11) generic ranking characters. Assessing the limits and potentials, the humility and hybris attitudes towards understanding reality, we hypothesise an integrative power of organisation, instead of simplistic natural selection, that works both in isolated and sympatric populations to maintain the integer state of the integrated autonomic entities against mutations, permutations, transmutations and perturbations. The permanent flux of these adverse, neutral or advantegous external and internal effects generates and gives rise to a responsive and balancing integrative mechanisms with comparative magnitude and multitude. This permanently balancing organisation process develops and drives a large number of interactions along various patterns of supervenient emergent mechanisms in order to integrate them into an oscillating autonomy. In isolation of the sky-islands on high altitudes, the divergences of new taxa are the adaptive autonomic byproducts of this integrative organisation. Integration and not the selection is the real force mechanism that harmonises the concerted flux of stochastic processes into reproductive isolation by sexual selection. The primacy of integration is clearly demonstrated by empirical evidences in the causes and consequences of the integrated speciation traits. Any kind of selection, artificial, natural, social or sexual, is only an emergent perturbating mechanism forced to integrate into the autonomy of an entity. This is how the entanglement of the quantum world and any supervenient integer work together through energeticism of the interactive realism. This is a must in the ontic, epistemic and semantic structural realism.

A description of the larva of Drusus plicatus Radovanović is given for the first time. The most important diagnostic characters enabling separation from larvae of the other Drusinae from the southeast Europe are listed. Molecular, ecological, and ethological features and distribution patterns of the species are given. Additionally, information on the sympatric caddisfly species of the three springs where larvae and adults of Drusus plicatus were found and presented.

The Balkan Peninsula is one of the most important European hotspots of freshwater biodiversity. The region is, however, to a large extent insufficiently investigated. Here we present data on distribution of caddisflies in one particularly understudied area, the Republic of Kosovo. Our data include the first records of Adicella altandroconia Botosaneanu & Novak and Halesus tessellatus (Rambur) for the Kosovo caddisfly fauna, and a new locality for the recently described Ecclisopteryx keroveci Previšić, Graf, & Vitecek. Further, we describe the new caddisfly species Drusus dardanicus sp. nov. from the Kopaonik Mountains. The new species belongs to the D. discophorus Species Group and differs morphologically from its most similar congeners (D. discophorus Radovanović, D. balcanicus Kumanski, and D. bureschi Kumanski) mainly in exhibiting (1) subtrianglar superior appendages ; (2) a narrow, dorsal spinate area of tergite VIII ; and (3) evenly rounded tips of intermediate appendages in caudal view. In phylogenetic analysis, D. dardanicus sp. nov. is well delineated and recovered as a sister taxon to D. osogovicus Kumanski, a species recorded from Bulgaria. The recent discovery of a new species and other rare or microendemic species presents important contributions to the knowledge on the rich freshwater biodiversity in Kosovo. These species face increasing anthropogenic pressure and threats to their conservation.

In this paper we describe Drusus sharrensis sp. n., from the Sharr Mountains in Kosovo. Males of the new species are morphologically most similar to Drusus krusniki Malicky, 1981, Drusus kerek Oláh, 2011 and Drusus juliae Oláh, 2011 but differ mainly in exhibiting (1) a differently shaped spinose area on tergite VIII; (2) intermediate appendages anteriorly curved in lateral view with broad tips in dorsal view; (3) inferior appendages with a distinct dorsal protrusion in the proximal half. Females of the new species are morphologically most similar to Drusus krusniki, Drusus kerek, Drusus juliae, and Drusus plicatus Radovanovic, 1942 but mainly differ in (1) segment X that is longer than the supragenital plate with distinctly pointed tips; (2) supragenital plate quadrangular with a distinct round dorsal protrusion; (3) a vulvar scale with a small median lobe. Results of phylogenetic species delimitation support monophyly of Drusus sharrensis sp. n. and recover it as sister to a clade comprising (Drusus pelasgus Oláh, 2010 + Drusus juliae + Drusus arbanios Oláh, 2010 + Drusus plicatus + (Drusus dacothracus Oláh, 2010 + Drusus illyricus Oláh, 2010)). The new species is a micro-endemic of the Sharr Mountains, a main biodiversity hotspot in the Balkan Peninsula. Main threats to the aquatic ecosystems of this part of the Balkan Peninsula are discussed.

In this study we present morphological, molecular and ecological features of the last instar larvae of Drusus bosnicus with data about distribution of this species in Bosnia and Herzegovina. We also included the most important diagnostic features enabling separation of larvae of D. bosnicus from larvae of the other European Drusinae and Trichoptera species.

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.

The Drusinae subfamily consists of about 100 species distributed across Europe. Some species are widespread and common, while others are restricted to one of several regional centers of endemism, such as in the Alps or the Balkan Highlands. These local and rare species, along with some newly described ones, are known from few confirmed occurrences only. While the full distribution of some species thus remains uncertain, the majority of species in the group are micro and regional endemics. Moreover, and because of the predominantly montane habitats they occupy, Drusinae are particularly susceptible to climate change. These aspects make this larger, running water taxon an interesting group to assess their vulnerability to environmental change through species distribution models (SDMs). Models are applied for individual Drusinae species to describe their current distribution patterns in Europe at a spatial resolution of 1 km. Beyond the usual application of bioclimatic predictors, additional predictors enhance the model: (i) topographical properties are used to approximate position in the catchment (catchment size) and (ii) soil properties indicate the possible composition of the substrate. Distribution predictions are projected for the year 2080 to determine changes in range size, as well as shifts in altitude and latitude/longitude. Projections are based on 5 different future climate models and 4 different future climate scenarios. Results of the first SDM models show mostly shifts towards northern latitudes and higher altitudes, for most of the future climate scenarios.

Adults and larvae of two new cryptic, endemic caddisflies, Ecclisopteryx keroveci sp.n. and Ecclisopteryx ivkae sp.n., are described and illustrated from the Western Balkans. Phylogenetic analysis (Bayesian MCMCMC) and association of different life history stages in both cryptic species were achieved through comparison of morphological characters and mitochondrial (mtCOI and mtLSU) and nuclear (nuWG) gene sequence data. The new species form a sister clade to the widely distributed E. dalecarlica and E. guttulata, with which they were formerly misidentified. Adults differ from each other and other species in the genus by the uniquely shaped inferior appendages in males and segment X in females. The larvae differ from each other and their congeners in the shape of the pronotum, and presence and constitution of additional spines on the parietalia. Larvae of both species are grazers and prefer stony substrate. Ecclisopteryx keroveci sp.n. has a wide distribution in the Western Balkans, while E. ivkae sp.n. is endemic to Dalmatia. Our findings demonstrate the significance of the Western Balkans as a freshwater biodiversity hotspot, and accentuate the importance of research focused on freshwater biodiversity and biogeography in southern Europe.

The caddisfly genus Drusus STEPHENS comprises roughly 100 species that inhabit mountain ranges in Europe, Asia Minor and the Caucasus. Here we present three new species of the genus Drusus and a hitherto unknown larva of Drusus macedonicus. All these species are closely related to Drusus discolor, Drusus romanicus romanicus and D. r. meridionalis, with which they share distinct features in feeding ecology, larval head capsule shape and adult morphology. The new species are clearly distinguished in both the larval and adult stages by unique features. Larvae can be identified by examination of frontoclypeal shape and surface structure. The adults can be separated from other species by details in the genital apparatus. Additionally all these species are clearly differentiated genetically from another and all previously known species of Drusus. Drusus krpachi n. sp., D. macedonicus, and D. sutjeskae n. sp. are micro-endemics of the western Balkan ranges ; D. bucegi n. sp. is an endemic of the Bucegi Mts in the south-eastern Carpathians.

Subfamily Drusinae is divided into 8 genera: Anomalopterygella, Cryptothrix, Drusus, Ecclisopteryx, Hadimina, Leptodrusus, Metanoea and Monocentra with about 100 described species. The genus with highest diversity is Drusus with 81 described species, whereas Anomalopterygella, Cryptothrix, Leptodrusus, Monocentra and Hadimina are monospecific gernera. Drusinae species usually inhabit springs and crenal sections of mountain streams and rivers with low temperatures and small annual amplitudes. Patchy distributions and isolation of populations are the reasons for the high diversity and high number of endemic species in subfamily Drusinae. Therefore, this subfamily is very interesting for taxonomic, phylogenetic, phylogeographic and evolutionary studies. This year we launched the project: “The Drusinae (Insecta: Trichoptera) in a world of global change: bridging basic and applied research in a highly sensitive aquatic insect group”. Coordinator of the project is Professor Johann Waringer from the University of Vienna. Trichopterologists from several countries are included in the project. They will perform collections in the field (larvae, pupae, adults, longterm water temperature data), descriptions of morphological features of undescribed larvae and undescribed taxa, DNA amplification and sequencing, phylogenetic analyses and species distribution modelling. The project has three main objectives. First, describing of morphological features of unknown larvae of 59 species from subfamily Drusinae and construction of identification keys. Second, to explore the phylogeny of Drusinae in a combined morphological and molecular context. We will test if key innovations (e.g., advanced feeding types) together with Pliocene– Pleistocene climatic change, promoted diversification and speciation. To test this hypothesis we need to increase the number of taxa included in our already established phylogeny of 28 species. We will include morphological data sets of larvae and adults as well as further molecular markers important for the reconstruction of Drusinae phylogeny. Third, analyses of developmental temperature data and species distribution modelling will provide valuable basic information for understanding high mountain biodiversity under two different future climate scenarios. This research project will continue in the next three years.

The Dinaric western Balkan karst area is a European biodiversity hotspot with particularly high degrees of endemism in the freshwater and subterranean fauna. For animals inhabiting surface waters karst development results in increased fragmentation of habitats. The reduction of surface waters and parallel development of higher energy relief may drive small scale vicariance of aquatic insect populations. We hypothesise that karstification mediated fragmentation of suitable surface water habitats for caddisflies and drove small scale vicariant diversification in the region. Using mtDNA sequence data, we apply a comparative phylogeographic approach to analyse population genetic structure of four ecologically similar Drusus species (Trichoptera, Limnephilidae) with different distribution ranges, date intra- and interspecific divergence of our target species, and link our findings with geological events. We observed marked differences in population genetic structure of our target species: three range-restricted endemics exhibited extremely high intraregional population differentiation dating to the late Pliocene/Pleistocene ; the widespread species had shallower divergence (Pleistocene) among regional clades of polyphyletic origin. These results suggest that the population histories of the endemic species differ from those of the more widespread species: populations of the endemic species differentiated within the study area due to regional fragmentation, potentially resulting from intensified karstification ; whereas the more widespread species repeatedly colonised the region from different sources.

According to our present knowledge, European benthic filter feeders have evolved only in the insect groups of Ephemeroptera, Trichoptera and Diptera. Caddisflies have refined these techniques by net-spinning as shown by most families of Hydropsychoidea (Hydropsychidae, Ecnomidae and Polycentropodidae) as well as in Philopotamidae ; in addition, genera Allogamus, Brachycentrus and Drusus developed extraordinary sets of filtering bristles at femora, heads and sometimes sterna, sometimes combined with elongations of femora or with strange head morphologies. Dipterans use nets like genus Rheotanytarsus or a modified labrum with retractable head fans as in Simuliidae. This presentation focuses on the importance of filtering species among netless larvae of Trichoptera within stream communities. Within Brachycentridae only genus Brachycentrus has evolved bristles originating from elongated femora for optimizing filtering while the second genus Micrasema feeds on mosses and fine detritus and consequently shows no adaptations at all. Most limnephilids are shredding omnivores (with the exception of some grazers like Melampophylax) but the genus Drusus has developed a remarkable radiation in feeding types (grazers, shredders and filter feeders). As a genus should represent both an ecotype and a morphotype, these findings have important taxonomic consequences. In contradiction to the River Continuum Concept netless filter feeders among caddiflies cover the crenal parts of high altitude mountains from 2000 m a.s.l. down to epi- and metapotamal lowland rivers at 200 m a.s.l. in high densities. This phenomenon seems to be a European peculiarity as no netless filtering species of caddisflies are known in North America except family Brachycentidae (Wiggins, 1996). In Asia the genus Limnocentropus takes the role of filtering Drusus species and occupies a similar niche at high-current microhabitats of torrents. Its general morphology shows strong analogies to Drusus and Cryptothrix.

A larval key of the Drusinae of Central Europe and the dinaric western Balkan is presented. Phylogeographic relationships are discussed in the light of molecular genetics, feeding ecology and larval morphology.

In this poster presentation, the fifth instar larva of Drusus vespertinus Marinković-Gospodnetić 1976 is described and illustrated. As most Drusinae, D. vespertinus is a cold-water tolerant species distributed in springs and crenal sections of the streams. It is one of seven species endemic to the Dinaric region that belong to the bosnicus group. All these species are range-restricted endemics, closely related and highly similar in adult and larval morphology. However, larvae of the majority of them have hitherto remained undescribed. Hence, here we give the most important diagnostic features that enable separation of D. vespertinus larva from larvae of the other species from the bosnicus group, but also from the other Drusinae. Overall, larva of D. vespertinus is distinguished from the other Drusinae by the presence of spinules (small spines) in the area around the eyes, by specific shape of pronotum and the presence of small white setae on dorsal surface of pronotum.