Research Article |
Corresponding author: Peter Manko ( peter.manko@unipo.sk ) Academic editor: Gunnar Kvifte
© 2021 Jan Ježek, Peter Manko, Jozef Oboňa.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Ježek J, Manko P, Oboňa J (2021) Psychodidae (Diptera) of Azerbaijan and Georgia – faunistics with biodiversity notes. ZooKeys 1049: 15-42. https://doi.org/10.3897/zookeys.1049.66063
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Records of 46 Psychodidae (Sycoracinae 1, Trichomyiinae 1, Psychodinae 44) species/subspecies are presented in this paper based on specimens collected by sweep-netting in Azerbaijan and Georgia in 2019. Nine species are recorded for the first time since their original description; 12 species are new for Transcaucasia; 22 species are new for Azerbaijan; and 17 species are new for Georgia. Saraiella ressli montana Ježek, 1990 is proposed as a synonym of S. ressli Wagner, 1983, syn. nov. Knowledge of some aspects of the ecology and biogeography of selected (especially rare) species has been expanded and a clear pattern was found in species richness, rare species, and new records in relation to land use, habitat diversity, and preservation of the environment surrounding the sampling site.
Biodiversity, Caucasus, faunistics, moth flies, new records, new synonymy, taxonomy, zoogeography
The purpose of faunistic studies is the registration of species. Intensive faunistic studies are necessary if we want to determine species richness on a local scale and track long‐term changes in species diversity (
Psychodidae is a relatively species-rich family, with nearly 500 species known to occur in Europe and the adjacent island areas; for more details, see
The Caucasus consists of several mountain ranges with different connectivity and serves heterogeneous environments, with different climatic conditions between lowlands and highlands as a result of orogenic activity in the Miocene-Pliocene (
Recent collecting sites on a map of the area of interest. Different colours (altitude) indicate the elevation in m a.s.l., and circle diameters and colours indicate the number of species recorded at each sampling site. The number of sites where the specific number of species is present is given in brackets.
Following our results and findings in Transcaucasia (
Dipterans were collected by PM, JO, T. Kovács (Mátra Museum of Hungarian Natural History Museum, Gyöngyös, Hungary), D. Murányi (Eszterházy Károly University, Eger, Hungary), and G. Vinçon (Grenoble, France) in the three periods between iv–v, vii, and ix–x, all in 2019, by sweep-netting from vegetation growing along watercourses and lakes. The captured specimens were preserved in 75% ethanol in the field. In the laboratory, specimens of Psychodidae were cleared in chloralphenol, treated in xylol, and mounted on glass slides in Canada balsam, subsequently identified by JJ and deposited at the National Museum (Natural History Museum), Department of Entomology, Prague, Czech Republic. The slides are labelled with the inventory slide number of the family Psychodidae (Inv. No.) and are included in the Diptera collection of National Museum Prague collections (
In the list of localities in Suppl. material
Abbreviations used:
TK Tibor Kovács;
DM Dávid Murányi;
GV Gilles Vinçon.
The distribution of the sampling sites (Fig.
We analysed material obtained in 2019 from 80 sites located in Transcaucasia in the territories of Azerbaijan and Georgia. Based on 182 slides from this material, we identified 46 species/subspecies belonging to three subfamilies. Below we present a list of recorded species and their distribution, together with the information on the material examined and notes.
Georgia: G 13, 8.7.2019 (Fig.
Occurrence of individual species on a map of the area of interest. Different colours (altitude) indicate the elevation in m a.s.l. and different symbols mark the sampling sites with occurrence of the species. When species co-occurred at the locality, an offset is shown in the legend and described in the Materials and methods. The number of sites where the species is present is given in brackets.
Caucasian species, not rare in Abkhazia (
Species known only from the original description from Abkhazia (
Azerbaijan: A 02, 9.5.2019, 1♂, slide Inv. No. 25663, leg. JO (Fig.
Rare species, known from Austria, Belgium, Czech Republic, Denmark, France, Germany, Great Britain, Greece, Hungary, Ireland, the Netherlands, Norway, Poland, Romania, Slovakia, and Sweden; larvae are xylophagous and occur in shaded slope spring areas and some other habitats with decaying organic matter (
First record for Azerbaijan and Transcaucasia.
Azerbaijan: A 14, 7.5.2019, 1♂, slide Inv. No. 25708, leg. JO; A 22, 5.5.2019, 1♂, slide Inv. No. 25625, leg. JO. Georgia: G 09, 2.5.2019, 2♂♂, slide Inv. No. 25642 and 25727, leg. JO; G 14, 2.5.2019, 1♂, slide Inv. No. 25710, leg. JO; G 26, 15.7.2019, 1♂, slide Inv. No. 25583, leg. TK, DM, GV; G 40, 13.7.2019, 1♂, slide Inv. No. 25633, leg. TK, PM, DM, GV (Fig.
Rare Caucasian species known only from a few localities (
Described from Georgia by
Georgia: G 03, 27.9.2019, 1♀, slide Inv. No. 25604, leg. PM (Fig.
Coprophagous species with a Western Palaearctic distribution. Known from Europe (Belgium, Great Britain, France, Germany, Ireland, the Netherlands, and Norway) (
First record for Georgia and Transcaucasia.
Georgia: G 09, 2.5.2019, 1♀, slide Inv. No. 25731, leg JO (Fig.
A common European species, distributed in Central Europe, along the Atlantic coast, in the British Isles and reaching the Apennines and the Balkans in the south, known from habitats with sprayed moss cushions, spring areas, and stream meanders across a wide range of elevations (
First record for Georgia and Transcaucasia.
Georgia: G 28, 6.7.2019, 1♂, slide Inv. No. 25609, leg. PM (Fig.
Generally common species, occurring across Europe to Transcaucasia, associated with fungi (
First record for Georgia, from a territory other than Abkhazia.
Azerbaijan: A 9, 6.5.2019, 1♀, slide Inv. No. 25716, leg. JO; A 18, 10.5.2019, 1♀, slide Inv. No. 25644, leg. JO; A 20, 5.5.2019, 1♀, slide Inv. No. 25720, leg. JO; A 21, 8.5.2019, 1♀, slide Inv. No. 25752, leg. JO; A 22, 5.5.2019, 1♀, slide Inv. No. 25627, leg. JO; A 23, 6.5.2019, 2♀♀, slide Inv. No. 25688 and 25695, leg. JO. Georgia: G 03, 27.9.2019, 1♀, slide Inv. No. 25605, leg. PM; G 18, 1.5.2019, 1♀, slide Inv. No. 25715, leg. JO; G 49, 28.4.2019, 1♀, slide Inv. No. 25670, leg. JO; G 56, 26.4.2019, 1♀, slide Inv. No. 25696, leg. JO (Fig.
A cosmopolitan species known from Europe (Austria, Azores, Belgium, Bosnia and Herzegovina, Bulgaria, Czech Republic, Denmark, Finland, France, Germany, Great Britain, Greece, Hungary, Ireland, Italy, Luxemburg, Madeira, the Netherlands, Norway, Poland, Portugal, Romania, Russia, Sardinia, Serbia, Slovakia, Slovenia, and Sweden). In Asia from Armenia, Azerbaijan, Afghanistan, China, India, Japan, North Korea, Syria, and Turkey. In Africa, from Algeria, the Canary Islands, Gambia, South Africa, and Tunisia. Also from Australia, New Zealand, South America, USA, Campbell Island, Juan Fernandez Islands, Kerguelen Island, and Macquarie Island (
First record for Georgia.
Azerbaijan: A 21, 8.5.2019, 1♀, slide Inv. No. 25751, leg. JO; A 23, 6.5.2019, 1♀, slide Inv. No. 25689, leg. JO (Fig.
Common Holarctic species, eurybiotic (
Azerbaijan: A 22, 5.5.2019, 1♀, slide Inv. No. 25626, leg. JO (Fig.
Uncommon Holarctic species. Recorded from Austria, Belgium, Czech Republic, France, Great Britain, Norway, Poland, Slovakia, Spain (incl. Madeira); Antipodes Is., Auckland L. (lake or lakes), Australia, Campbell L., Chile, Enderby L., Ewing L., French L., Macquarie L., New Zealand, Ocean L., Rose L., Saint Helena, and USA (
First record for Azerbaijan and Transcaucasia.
Georgia: G 23, 29.4.2019, 1♀, slide Inv. No. 25714, leg. JO (Fig.
Common European and Western Siberian species recorded from Belgium, Czech Republic, Greit Britain, Slovakia, Slovenia, and the Netherlends (
First record for Georgia and Transcaucasia.
Azerbaijan: A 14, 7.5.2019, 3♂♂, slide Inv. No. 25738, 25739 and 25740, leg. JO (Fig.
Species known only from the original description from Abkhazia (
Azerbaijan: A 02, 9.5.2019, 1♂, slide Inv. No. 25661, leg. JO (Fig.
Species known only from Czech Republic, Greece, Slovakia, and Slovenia; until now considered as a rare European species (
Based on the record from the southern slope of the Greater Caucasus foothills (see above), it is necessary to reconsider it as a European and Transcaucasian species. First record for Azerbaijan and Transcaucasia.
Azerbaijan: A 23, 6.5.2019, 1♂, slide Inv. No. 25693, leg. JO (Fig.
Locally common species, known from several European countries: Belgium, Austria, Great Britain, Ireland, Bosnia and Herzegovina, Czech Republic, Greece, and Macedonia (
Records scattered over a large area of Transcaucasia (Fig.
Azerbaijan: A 19, 7.5.2019, 1♂, 1♀, slide Inv. No. 25671 and 25723, leg. JO. Georgia: G 19, 30.4.2019, 1♂, slide Inv. No. 25649, leg. JO; G 21, 3.5.2019, 1♀, slide Inv. No. 25681, leg. JO (Fig.
Holarctic species, known from most of Europe, very common locally, mainly in extreme localities (salt works, thermal springs, calcareous water, mineral-rich springs). Also recorded in Algeria, Morocco, Israel, Armenia, Azerbaijan, Afghanistan, Iran, and USA (
First record for Georgia.
Georgia: G 24, 29.4.2019, 3♂♂, slide Inv. No. 25678, 25734 and 25745, leg. JO; G 35, 11.7.2019, 1♂, slide Inv. No. 25616, leg. TK, PM, DM; G 34, 11.7.2019, 1♂, slide Inv. No. 25581, leg. TK, PM, DM; G 35, 11.7.2019, 1♂, slide Inv. No. 25571, leg. TK, PM, DM, GV; G 43, 11.7.2019, 1♂, slide Inv. No. 25617, leg. TK, PM, DM, GV (Fig.
Bazarella joosti Vaillant, 1983, was described from Schelda nr. Kashadl (Central Caucasus: Georgia, probably Gora Shkhel’da, omitted (overlooked) from Georgia in
Parabazarella Vaillant, 1983 was raised to genus level by
Azerbaijan: A 01 9.5.2019, 1♂, slide Inv. No. 25624, leg. JO; A 02, 9.5.2019, 1♂, slide Inv. No. 25665, leg. JO; A 23, 6.5.2019, 1♂, slide Inv. No. 25687, leg. JO. Georgia: G 02, 24.9.2019, 1♂, slide Inv. No. 25584, leg. TK, PM; G 09, 2.5.2019, 1♂, slide Inv. No. 25732, leg. JO; G 25, 29.4.2019, 1♂, slide Inv. No. 25713, leg. JO; G 47, 9.7.2019, 1♂, slide Inv. No. 25638, leg. TK, PM, DM, GV.
Common European and Transcaucasian species, penetrates into North Africa (
Azerbaijan: A 01, 9.5.2019, 1♂, slide Inv. No. 25707, leg. JO; A 02, 9.5.2019, 1♂, slide Inv. No. 25664, leg. JO; A 13, 7.5.2019, 1♂, slide Inv. No. 25743, leg. JO; A 14, 7.5.2019, 1♂, slide Inv. No. 25709, leg. JO. Georgia: G 20, 3.5.2019, 1♂, slide Inv. No. 25717, leg. JO; G 24, 29.4.2019, 1♂, slide Inv. No. 25749, leg. JO.
European, western Siberian, and Caucasian species (
First record for Georgia, from a territory other than Abkhazia.
Azerbaijan: A 02, 9.5.2019, 1♂, slide Inv. No. 25662, leg. JO; A 13, 7.5.2019, 1♂, slide Inv. No. 25742, leg. JO. Georgia: G 24, 29.4.2019, 1♂, slide Inv. No. 25747, leg. JO.
Not a common European species, known from Czech Republic, Denmark, Finland, France, Norway, Slovakia, and Ukraine (
Occurrence of individual species on a map of the area of interest. Different colours (altitude) indicate the elevation in m a.s.l. and different symbols the sampling sites with occurrence of the species. When species co-occurred at a locality, an offset is shown in the legend and described in the Materials and methods. The number of sites where the species is present is given in brackets.
It is necessary to reconsider this as a European and Transcaucasian species. First record for Azerbaijan, Georgia, and Transcaucasia.
Azerbaijan: A 04, 1.10.2019, 1♂, slide Inv. No. 25597, leg. PM; A 05, 1.10.2019, 1♂, slide Inv. No. 25578, leg. PM; A 06, 1.10.2019, 1♂, slide Inv. No. 25574, leg. PM. Georgia: G 01, 25.9.2019, 1♂, slide Inv. No. 25601, leg. PM; G 41, 27.4.2019, 2♂♂, slides Inv. No. 25653 and 25654, leg. JO.
Species known from Bosnia and Herzegovina, Bulgaria, Montenegro, Serbia, Macedonia, Armenia, and Azerbaijan (
First record for Georgia.
Georgia: G 17, 30.4.2019, 1♂, slide Inv. No. 25657, leg. JO; G 24, 29.4.2019, 2♂♂, slide Inv. No. 25736 and 25746, leg. JO; G 41, 27.4.2019, 1♂, slide Inv. No. 25652, leg. JO.
Species widespread in Europe, North Africa, and Transcaucasia (Armenia, Azerbaijan). In Europe, known from Albania, Austria, Belgium, Bosnia and Herzegovina, Bulgaria, Czech Republic, Crete, Croatia, France, Germany, Great Britain, Greece, Hungary, Ireland, Italy, Macedonia, Montenegro, Poland, Serbia, Slovakia, Slovenia, Spain, and Ukraine (
First record for Georgia.
Azerbaijan: A 05, 1.10.2019, 1♂, slide Inv. No. 25577, leg. PM.
Species known only from the original description from Greece (Vaillant 1978).
Extremely rare species. First record since the original description, and therefore a first record for Azerbaijan.
Azerbaijan: A 04, 1.10.2019, 1♂, slide Inv. No. 25596, leg. PM.
Distribution. An extremely rare species known from Bulgaria, Georgia, Greece, Macedonia, Serbia, and Romania (
First record for Azerbaijan.
Georgia: G 11, 3.5.2019, 1♂, slide Inv. No. 25674, leg. JO; G 16, 2.5.2019, 1♂, slide Inv. No. 25680, leg. JO.
Species known from the whole of Europe: Austria, Belgium, Bulgaria, Czech Republic, France, Germany, Greece, Great Britain, Hungary, Ireland, Italy, Slovakia, Slovenia, Spain, and Switzerland (
First record for Georgia and Transcaucasia.
Azerbaijan: A 03, 1.10.2019, 1♂, slide Inv. No. 25588, leg. PM. Georgia: G 12, 9.7.2019, 1♂, slide Inv. No. 25612, leg. PM, GV.
This is a common European and western Siberian species. In Europe it is known from Austria, Belgium, Great Britain, Czech Republic, Denmark, European Turkey, France, Germany, Greece, Hungary, Slovakia, Sweden, and the Netherlands. In Asia, it is known from Armenia, Turkey, Kyrgyzstan, Afghanistan, and China; it occurs from lowlands to mountains, usually associated with mosses in running water habitats; its larvae are rheobiotic (
Occurrence of individual species on a map of the area of interest. Different colours (altitude) indicate the elevation in m a.s.l. and different symbols the sampling sites with occurrence of the species. When the species co-occurred at the locality, an offset is shown in the legend and described in the Materials and methods. The number of sites where the species is present is given in brackets.
First record for Azerbaijan and Georgia.
Georgia: G 02, 24.9.2019, 1♂, slide Inv. No. 25585, leg. TK, PM.
The nominal subspecies was recorded from several European countries and is also known from Abkhazia in the Transcaucasia; it lives in forest slope spring areas, brooks, and marshes (
First records for Georgia outside of Abkhazia.
Azerbaijan: A 02, 9.5.2019, 1♂, slide Inv. No. 25659, leg. JO; A 07, 30.9.2019, 1♂, slide Inv. No. 25590, leg. PM; A 08, 30.9.2019, 1♂, slide Inv. No. 25602, leg. PM; A 13, 7.5.2019, 1♂, slide Inv. No. 25741, leg. JO; A 21, 8.5.2019, 1♂, slide Inv. No. 25750, leg. JO; A 23, 6.5.2019, 1♂, slide Inv. No. 25684, leg. JO. Georgia: G 09, 2.5.2019, 1♂, slide Inv. No. 25641, leg. JO; G 25, 29.4.2019, 1♂, slide Inv. No. 25712, leg. JO.
This is a very common species which is recorded from throughout Europe, Armenia, and the Canary Islands. In Europe, it is known from Austria, Belgium, Bosnia and Herzegovina, Bulgaria, Croatia, Czech Republic, Denmark, Finland, France, Georgia, Germany, Great Britain, Greece, Hungary, Ireland, Italy, Luxembourg, Macedonia, Montenegro, Poland, Romania, Sardinia, Serbia, Slovakia, Slovenia, Spain, Switzerland, the Netherlands, and Ukraine; abundant especially in shaded habitats with decaying organic matter, e.g., ponds, brooks, spring areas, water reservoirs, and swamps (
First record for Azerbaijan.
Georgia: G 24, 29.4.2019, 1♂, slide Inv. No. 25677, leg. JO; G 48, 28.4.2019, 1♂, slide Inv. No. 25646, leg. JO.
European species, also registered from Turkey, Georgia, and the Canary Islands (
Azerbaijan: A 03, 2.10.2019, 1♂, slide Inv. No. 25587, leg. PM; A 04, 1.10.2019, 1♂, slide Inv. No. 25593, leg. PM; A 09, 30.9.2019, 1♂, slide Inv. No. 25582, leg. PM; A 10, 6.5.2019, 2♂♂, slide Inv. No. 25647 and 25675, leg. JO; A 12, 8.5.2019, 1♂, slide Inv. No. 25698, leg. JO; A 20, 5.5.2019, 1♂, slide Inv. No. 25721, leg. JO. Georgia: G 16, 2.5.2019, 1♂, slide Inv. No. 25679, leg. JO; G 29, 30.9.2019, 1♂, slide Inv. No. 25600, leg. GV; G 36, 27.4.2019, 1♂, slide Inv. No. 25667, leg. JO; G 39, 9.7.2019, 1♂, slide Inv. No. 25705, leg. PM; G 42, 4.7.2019, 1♂, slide Inv. No. 25620, leg. PM; G 44, 10.7.2019, 1♂, slide Inv. No. 25628, leg. TK, PM, DM; G 45, 10.7.2019, 1♂, slide Inv. No. 25607, leg. TK, PM, DM. JO; G 46, 1.10.2019, 1♂, slide Inv. No. 25608, leg. GV; G 49, 28.4.2019, 1♂, slide Inv. No. 25669, leg. JO; G 50, 28.4.2019, 1♂, slide Inv. No. 25656, leg. JO.
European species. Known also from Tajikistan, Azerbaijan, and Georgia (
Azerbaijan: A 21, 8.5.2019, 1♂, slide Inv. No. 25725, leg. JO. Georgia: G 24, 29.4.2019, 1♂, slide Inv. No. 25744, leg. JO.
A very common European species. Known from Austria, Belgium, Bosnia and Herzegovina, Croatia, Czech Republic, Denmark, Finland, France, Germany, Great Britain, Hungary, Ireland, the Netherlands, Norway, Poland, Serbia, Slovakia, Slovenia, Spain, Sweden, Switzerland, and Ukraine, in both shaded and unshaded habitats with decaying organic matter (ponds, brooks, spring areas, swamps, and water reservoirs) where larvae develop (
Occurrence of individual species on a map of the area of interest. Different colours (altitude) indicate the elevation in m a.s.l. and different symbols the sampling sites with occurrence of species. When species co-occurred at the locality, an offset is shown in the legend and described in the Materials and methods. The number of sites where the species is present is given in brackets.
First record for Azerbaijan, Georgia, and Transcaucasia.
Azerbaijan: A 23, 6.5.2019, 1♂, slide Inv. No. 25691, leg. JO.
Species known throughout Europe, including northern areas (British Isles, Scandinavian bioregion) and Central European countries, penetrating eastwards as far as Turkey (Anatolia) and Azerbaijan; southern frontier of distribution is limited by North Africa (
Azerbaijan: A 02, 9.5.2019, 1♂, slide Inv. No. 25666, leg. JO.
Widespread Holarctic polyvoltine species occurring from lowlands to mountains; known also from Georgia (e.g.,
Occurrence of individual species on a map of the area of interest. Different colours (altitude) indicate the elevation in m a.s.l. and different symbols the sampling sites with occurrence of the species. The number of sites where the species is present is given in brackets after the species name. When species co-occurred at the locality, an offset is also shown in the legend and described in the Materials and methods.
First record for Azerbaijan.
Azerbaijan: A 20, 5.5.2019, 1♂, slide Inv. No. 25722, leg. JO; A 23, 6.5.2019, 2♀♀, slide Inv. No. 25686 and 25694, leg. JO. Georgia: G 10, 4.5.2019, 1♀, slide Inv. No. 25711, leg. JO.
A very common cosmopolitan species ranging from lowlands to mountains. The larvae are saprobiotic, occasionally associated with fungi; the adults are often found in unclean bathrooms. Known from Austria, Azores, Belgium, Bosnia and Herzegovina, Bulgaria, the Canary Islands, Cyprus, Czech Republic, Denmark, Finland, France, Germany, Great Britain, Greece, Hungary, Ireland, Italy (incl. Sardinia), Madeira, the Netherlands, Norway, Poland, Romania, Russia, Serbia, Slovakia, Slovenia, Spain, Sweden, Switzerland, Ukraine, Turkey, Abkhazia, Afghanistan, Africa mer., Algeria, Argentina, Australia, Azores, Brazil, Canada, Chile, Iran, Israel, Juan Fernandéz Islands, New Zealand, Puerto Rico Islands, Tunisia, USA (
First record for Azerbaijan. First record for Georgia, from a territory other than Abkhazia.
Azerbaijan: A 05, 1.10.2019, 1♀, slide Inv. No. 25579, leg. PM; A 23, 6.5.2019, 1♀, slide Inv. No. 25685, leg. JO. Georgia: G 09, 2.5.2019, 1♀, slide Inv. No. 25733, leg. JO; G 19, 30.4.2019, 1♀, slide Inv. No. 25651, leg. JO; G 24, 29.4.2019, 1♂, 1♀, slide Inv. No. 25735 and 25748, leg. JO.
A common European species, it occurs commonly from lowlands to mountains. The larvae are saprobiotic and often develop in nests of water birds. Recorded from Austria, Belgium, Bosnia and Herzegovina, Croatia, Czech Republic, Great Britain, Denmark, France, Finland, Germany, Greece, Hungary, Ireland, Norway, Romania, Serbia, Slovakia, Slovenia, Spain, Switzerland, the Netherlands, Ukraine, and Georgia: Abkhazia (
First record for Azerbaijan.
Azerbaijan: A 23, 6.5.2019, 1♂, slide Inv. No. 25690, leg. JO; Georgia: G 05, 25.9.2019, 1♂, slide Inv. No. 25589, leg. PM.
A rare species associated with fungi, occurs from lowlands to mountains, so far known only from the Czech Republic, France, Slovakia, Slovenia, Switzerland, and Ukraine (
First record for Azerbaijan, Georgia, and Transcaucasia.
Azerbaijan: A 19, 7.5.2019, 1♂, slide Inv. No. 25724, leg. JO. Georgia: G 22, 15.7.2019, 1♂, slide Inv. No. 25599, leg. TK, DM, GV.
A very common Holarctic species. Known from Austria, Belgium, Bulgaria, Croatia, Czech Republic, Denmark, Finland, France, Germany, Great Britain, the Greek mainland, Hungary, Ireland, Italy, the Netherlands, Norway, Poland, Romania, Russia, Sardinia, Slovakia, Slovenia, the Spanish mainland, Sweden, Ukraine, Georgia, Algeria, and USA (
First record for Azerbaijan. It is a vector of larval stages of Rhabditis nematodes and Gamasidae mites (
S. ressli montana Ježek, 1990, syn. nov.
Azerbaijan: A 02, 9.5.2019, 1♂, slide Inv. No. 25660, leg. JO; A 04, 1.10.2019, 1♂, slide Inv. No. 25595, leg. PM; A 05, 1.10.2019, 1♂, slide Inv. No. 25576, leg. PM; A 06, 1.10.2019, slide Inv. No. 25573, 1♂, leg. JJ, PM; A 10, 6.5.2019, 2♂♂, slide Inv. No. 25676 and 25648, leg. JO; A 14, 7.5.2019, 1♂, slide Inv. No. 25737, leg. JO; A 16, 10.5.2019, 1♂, slide Inv. No. 25643, leg. JO; A 17, 10.5.2019, 1♂, slide Inv. No. 25700, leg. JO; A 18, 10.5.2019, 1♂, slide Inv. No. 25645, leg. JO; A 23, 6.5.2019, 1♂, slide Inv. No. 25692, leg. JO; A 24, 9.5.2019, 1♂, slide Inv. No. 25703, leg. JO. Georgia: G 03, 27.9.2019, 1♂, slide Inv. No. 25606, leg. PM; G 04, 27.9.2019, 1♂, slide Inv. No. 25592, leg. PM; G 09, 2.5.2019, 2♂♂, slides Inv. No. 25640, 25726, leg. JO; G 12, 9.7.2019, 1♂, slide Inv. No. 25614, leg. PM, GV; G 14, 2.5.2019, 1♂, slide Inv. No. 25658, leg. JO; G 15, 2.5.2019, 2♂♂, slides Inv. No. 25673, 25702 leg. JO; G 16, 28.9.2019, 1♂, slide Inv. No. 25586, leg. GV; G 31, 27.4.2019, 1♂, slide Inv. No. 25697, leg. JO; G 37, 2.10.2019, 1♂, slide Inv. No. 25701, leg. PM; G 38, 27.4.2019, 1♂, slide Inv. No. 25704, leg. JO; G 40, 13.7.2019, 1♂, slide Inv. No. 25632, leg.TK, PM, DM, GV; G 47; 9.7.2019, 1♂, slide Inv. No. 25636, leg. TK, PM, DM, GV; G 49, 28.4.2019, 1♂, slide Inv. No. 25668, leg. JO; G 50, 28.4.2019, 1♂, slide Inv. No. 25655, leg. JO; G 51, 28.4.2019, 1♂, slide Inv. No. 25699, leg. JO; G 52, 11.7.2019, 1♂, slide Inv. No. 25618, leg. TK, PM, DM, GV; G 53, 9.7.2019, 1♂, slide Inv. No. 25623, leg. TK, PM, DM, GV; G 54, 27.4.2019, 1♂, slide Inv. No. 25672, leg. JO; G 55, 29.9.2019, 1♂, slide Inv. No. 25580, leg. PM.
Species known from Iran, Armenia, Azerbaijan, and Russia (Wagner and Joost 1983;
Saraiella ressli Wagner, 1981, was described only on the basis of a holotype from the environment of the Caspian Sea (northern Iran, Veyshar nr. Nowshahr env. Chalus – Mazandaran, 1400 m a.s.l.). Saraiella ressli montana Ježek, 1990, was described from S. E. Iran, Kerman province, Kuh-e-Lalehzar, 3850–4374 m a.s.l. with top 4374 m (holotype and two paratypes), and additionally two paratypes from E. Iran, Kerman province, Deh Bakri, Kuh-e Jebal Barez (Jebal Barez Mts). The numerous materials from Caucasus (males) cited above proved a large variability of specimens (last three flagellomeres almost globular or long-oval, terminal flagellomere with small apiculus or without a protuberance, coxopodites basally with or without several bristles, gonostylus with a small subapical tooth terminally or not; all characters are mixed on the same localities). The subspecific rank is therefore groundless and S. ressli montana Ježek, 1990 is thus sunk as a synonym herein.
Azerbaijan: A 04, 1.10.2019, 1♂, slide Inv. No. 25594, leg. PM, A 06, 1.10.2019, 1♂, slide Inv. No. 25572, leg. PM. Georgia: G 07, 17.7.2019, 1♂, slide Inv. No. 25591, leg. GV.
European species, known from Bosnia and Herzegovina, Czech Republic, Italy, Poland, Serbia, and Slovakia (
First record for Azerbaijan, Georgia, and Transcaucasia.
Georgia: G 09, 2.5.2019, 1♂, slide Inv. No. 25729, leg. JO.
Species known only from the original paper from Abkhazia, where only one specimen was collected on the bank of a stream in 2000 m a.s.l. (
First record since the original description. First record for Georgia, a territory other than Abkhazia. An extremely rare species known from only two sites in Greater Caucasus.
Azerbaijan: A 05, 1.10.2019, 1♂, slide Inv. No. 25575, leg. PM.
Species known only from the original paper from Armenia (
First record since the original description, first record for Azerbaijan. An extremely rare species known from only two sites in the Lesser Caucasus at elevations higher than 1000 m.
Georgia: G 09, 2.5.2019, 1♂, slide Inv. No. 25728, leg. JO; G 30, 13.7.2019, 1♂, slide Inv. No. 25611, leg. TK, PM, DM, GV.
Species known from Austria, Bosnia, Czech Republic, France, Germany, Kosovo, Poland, Slovakia, Switzerland, and the United Kingdom as well as Abkhazia (
First record for Georgia, territory other than Abkhazia.
Georgia: G 06, 24.9.2019, 1♂, slide Inv. No. 25598, leg. GV.
Known only from the original paper from Abkhazia (
First record since the original description. First record for Georgia, from a territory other than Abkhazia. An extremely rare species known from only two sites at a distance ca. 20–30 km from the Black Sea coast at elevations of ca. 2000 and 2600 m.
Georgia: G 03, 27.9.2019, 1♀, slide Inv. No. 25603, leg. PM; G 08, 1.5.2019, 1♀, slide Inv. No. 25683, leg. JO; g 56, 26.4.2019, 1♀, slide Inv. No. 25682, leg. JO.
Cosmopolitan and euryvalent species (
Azerbaijan: A 17, 10.5.2019, 2♂♂, slide Inv. No. 25718 and 25719, leg. JO. Georgia: G 09, 2.5.2019, 1♂, slide Inv. No. 25730, leg. JO; G 40, 13.7.2019, 1♂, slide Inv. No. 25635, leg. TK, PM, DM, GV; G 42, 4.7.2019, 1♂, slide Inv. No. 25622, leg. PM; G 47, 9.7.2019, 1♂, slide Inv. No. 25637, leg. TK, PM, DM, GV.
Species known only from the original paper from Abkhazia (
First record since the original description, first record for Azerbaijan. Occurs at a wide range of elevations (810–3050 m a.s.l.) at small forest or bushy streams, springs, and side brooks of the Greater Caucasus.
Georgia: G 27, 17.7.2019, 1♂, slide Inv. No. 25570, leg. GV; G 33, 5.7.2019, 1♂, slide Inv. No. 25629, leg. PM, GV; G 40, 13.7.2019, 1♂, slide Inv. No. 25631, leg. TK, PM, DM, GV; G 42, 4.7.2019, 2♂♂, slide Inv. No. 25619 and 25621, leg. PM; G 47, 9.7.2019, 1♂, slide Inv. No. 25639, leg. TK, PM, DM, GV.
This is a common European species known from Austria, Czech Republic, Finland, France, Germany, Great Britain, Italy, Lithuania, Poland, Slovakia, Slovenia, Ukraine, and Armenia (
First record for Georgia. In Europe, a very common species from lowlands to mountains, but in Georgia it has been found only in mountain localities at an elevation of 2050 to 3050 m a.s.l.
Azerbaijan: A 15, 8.5.2019, 1♂, slide Inv. No. 25706, leg. JO. Georgia: G 19, 30.4.2019, 1♂, slide Inv. No. 25650, leg. JO.
Species known only from the original paper from Abkhazia (
First records since the original description, first record for Azerbaijan and Georgia, from a territory other than Abkhazia. An extremely rare species known from only three considerably remote localities in lowland and foothills of the Greater Caucasus (recent records at 655 and 845 m a.s.l.).
Although faunistic research of the family Psychodidae has been the subject of a number of authors (see the Introduction for references) and a large number of sites were examined in our recent studies (
One of our most interesting results is that we confirmed several species that were described from the Abkhazia region, and no one has been able to confirm them anywhere else until this study was carried out, despite several sampling campaigns. These are species that are extremely rare or relatively common in Abkhazia. Extremely rare was Seoda svanetica (
Based on our new data, knowledge of some aspects of the ecology and biogeography of selected (especially rare) species has also expanded. Our records point out that several rare species show different patterns related to elevation: Clytocerus (Boreoclytocerus) grusinicus and Tonnoiriella arcuata occur at large range of elevations, Mormia ckvitariorum and Yomormia afonensis prefer low elevations in lowlands and low foothills, Thornburghiella veve elevations higher than 1000 m, and Threticus petrosus sites at elevations higher than 2000 m relatively close to the Black Sea coast. Several species considered as European (or rare European) occur at numerous localities in the Transcaucasia and should be reconsidered as European and Transcaucasian species (Promormia silesiensis, Panimerus denticulatus, Pericoma (Pachypericoma) nielseni).
Although we usually found only one to three species in most sites located in cultivated and managed areas, mosaic croplands and mostly open areas with dominant herbaceous cover, the species number was quite high at other sampling sites. The highest species richness was recorded at sites A 23 (Fig.
The collecting site with the highest species richness in Azerbaijan (A 23), a karst brook in deciduous forest in the Şəki district, Şəki, Quirxbulaq, with seven species collected; general view (left) and different habitats (top right and bottom right which also include colleagues Libor Dvořák (left) and Ľuboš Hrivniak (right) collecting insects during a joint sampling campaign); photograph P. Manko.
Out of the total number of 46 species/subspecies (Sycoracinae 1 sp., Trichomyiinae 1 sp. Psychodinae 44 sp.) and 182 slides, 12 species are recorded for the first time for Transcaucasia (namely Copropsychoda brevicornis, Feuerborniella obscura, Logima sigma, L. zetterstedti, Panimerus denticulatus, Pericoma (Pachypericoma) nielseni, P. (Pericoma) pseudoexquisita, Pneumia trivialis, Promormia silesiensis, Psychomora mycophila, Saraiella rotunda, Trichomyia urbica), 22 species for Azerbaijan (namely Clytocerus (Boreoclytocerus) grusinicus, Logima sigma, Mormia ckvitariorum, Panimerus denticulatus, Pericoma (Pachypericoma) nielseni, Pericoma (Pericoma.) kariana, P. (P.) motasi motasi, Pneumia canescens, P. nubila, P. trivialis, Promormia silesiensis, Psychoda phalaenoides, Psychodocha cinerea, P. gemina, Psychomora mycophila, P. trinodulosa, Saraiella ressli, S. rotunda, Thornburghiella veve, Tonnoiriella arcuata, Trichomyia urbica, Yomormia afonensis), 17 species for Georgia (Copropsychoda brevicornis, Feuerborniella obscura, Logima albipennis, L. zetterstedti, Panimerus denticulatus, Parabazarella joosti lalehzarica, Paramormia (Duckhousiella) ustulata, Pericoma (Pachypericoma) nielseni, Pericoma (Pericoma) bosniaca, P. (P.) exquisita, P. (P.) pseudoexquisita, Pneumia canescens, P. trivialis, Psychomora mycophila, Saraiella ressli, S. rotunda, Ulomyia cognata), and nine species for the first time since their original descriptions (Clytocerus (Boreoclytocerus) grusinicus, Mormia ckvitariorum, Parabazarella joosti lalehzarica, Pericoma (P.) kariana, Sycorax caucasica, Thornburghiella veve, Threticus petrosus, Tonnoiriella arcuata, Yomormia afonensis). The knowledge of some aspects of ecology and biogeography of selected (especially rare) species has been expanded and a clear pattern was found in species richness, rare species, and new records in relation to land use, habitat diversity and preservation of the environment surrounding the sampling sites.
We are grateful to B. Japoshvili and L. Mumladze (Ilia State University, Tbilisi, Georgia) and N. Snegovaya (Azerbaijan National Academy of Sciences, Baku, Azerbaijan) for their kindness and help during the fieldwork and for providing permits for the collection and export of material. Thanks are due mainly to T. Kovács (Mátra Museum of Hungarian Natural History Museum, Gyöngyös, Hungary), D. Murányi (Eszterházy Károly University, Eger, Hungary) and G. Vinçon (Grenoble, France) for providing extensive additional material from the Caucasus. The authors thank Štefan Koco (University of Prešov, Slovakia) for advice in preparing the maps and providing additional information on land use in the wider area of the collecting sites. We would also like to thank Gunnar Kvifte (Steinkjer, Norway), Alessio Morelli (Pianella, Italy) and Nathalie Yonow (Swansea, United Kingdom) very much for contributing to the improvement of the manuscript with their comments and corrections. This work was supported by the Ministry of Culture of the Czech Republic (DKRVO 2019–2023/5.I.c, National Museum, 00023272) and by the Cultural and Educational Grant Agency (The Ministry of Education, Science, Research and Sport of the Slovak Republic) under contract No. 005PU-4/2019.
Table S1. List of localities with altitudes and coordinates
Data type: occurrence