Psychodidae (Diptera) of Azerbaijan and Georgia – faunistics with biodiversity notes

Abstract 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.


Introduction
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 longterm changes in species diversity (Ejsmont-Karabin 2019). This applies in particular to areas that are of the utmost importance from the point of view of biodiversity but are still insufficiently researched for some groups of organisms. One such biodiversity hotspot is the Caucasus, and one of its hitherto insufficiently known groups is the family Psychodidae. For the above reasons, this is another contribution to knowledge of the Diptera: Psychodidae, especially Sycoracinae, Trichomyiinae and Psychodinae from Azerbaijan and Georgia.
Following our results and findings in Transcaucasia (Ježek et al. 2018(Ježek et al. , 2021Oboňa et al. 2017Oboňa et al. , 2019a, which indicate the relatively high diversity of this family in the Caucasus, we continued to collect material in 2019. We did so in an effort to better understand the diversity and occurrence of the family Psychodidae in the Caucasus and to make the data on the faunistics and biodiversity of this group more complete.

Materials and methods
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 (NMPC), see Tkoč et al. (2014). The nomenclature is modified from Vaillant (1972Vaillant ( -1983 and Wagner (1990Wagner ( , 2018 using the classifications of Ježek (1990aJežek ( , 2007; Ježek and van Harten (2005); Omelkova and Ježek (2012); Oboňa and Ježek (2014) and Ježek (2015, 2019). We use the term "Transcaucasia" (Armenia, Azerbaijan, and Georgia) according to the Catalogue of Palaearctic Diptera; for more details, see Wagner (1990). Information on the distribution (simple distribution overview) is given for the species recorded from Azerbaijan and Georgia for the first time.
In the list of localities in Suppl. material 1: Table S1, data are presented in the following order: number (in brackets), territorial unit (district, region etc.), name of the locality, nearest settlement (where appropriate), specified location and habitat, coordinates, and elevation. The localities are listed alphabetically. Data for the material examined are arranged in the following order: number, date of collection, number of males and females, slide number and collector's or determinator's name (abbreviated).
Abbreviations used: The distribution of the sampling sites ( Fig. 1) and species recorded in this study (Figs 2-7) are presented in maps prepared using data derived from USGS/NASA SRTM data providing seamless continuous topography surfaces (Jarvis et al. 2008). Areas with different elevations were painted in QGIS (version: 3.10.10-A Coruña) with the 'Band Rendering' (Singleband Pseudocolor, Oranges Colour Ramp, the colour for values from > 0 to ≤ 500 was changed to green). When species co-occurred at the locality, the offset was set to x = 1 mm, y = 1 mm for species marked with square symbol, x = -1 mm, y = -1 mm for species marked with triangles and x = 1 mm, y = -1 mm for diamonds, respectively.

Results
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.

Subfamily Sycoracinae
Sycorax caucasica Ježek, 1990 Material examined. Georgia: G 13, 8.7.2019 ( Fig. 2), 1♂, slide Inv. No. 25630, leg. PM, GV. Distribution. Caucasian species, not rare in Abkhazia (Ježek 1990a). Note. Species known only from the original description from Abkhazia (Ježek 1990a). First record since the original description, first record from Georgia, from a territory other than Abkhazia. Although this is a typical mountain species (elevation of ca. 2000 m a.s.l.), it is also known from the small hills (Ježek 1990a) and foothills of the Greater Caucasus (see above).
Distribution. 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 (Ježek 2003;Ježek and Omelková 2012).
Note. First record for Azerbaijan and Transcaucasia.

Distribution.
Rare Caucasian species known only from a few localities (Wagner 1981;Oboňa et al. 2019).
Note. Described from Georgia by Wagner (1981) on the basis of the only holotype from a right tributary above the village Kwarchi. After the second record from Georgia (Oboňa et al. 2019), the species was recorded from another six sites in this study. Species found in low numbers in a range of elevations from 295 m (Oboňa et al. 2019) to 1520 m a.s.l. (see above). First records for Azerbaijan.
Note. First record for Georgia and Transcaucasia.

Feuerborniella obscura (Tonnoir, 1919)
Material examined. Georgia: G 09, 2.5.2019, 1♀, slide Inv. No. 25731, leg JO (Fig. 2). Distribution. 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, 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. and stream meanders across a wide range of elevations Oboňa et al. 2019b).
Note. First record for Georgia and Transcaucasia.
Note. First record for Georgia, from a territory other than Abkhazia.  Ježek et al. 2018b).
Note. Records scattered over a large area of Transcaucasia ( Fig. 3) suggest that the species is also relatively common in Transcaucasia. It is necessary to reconsider it as a European and Transcaucasian species. First record for Azerbaijan, Georgia, and Transcaucasia.  (Fig. 3).
Note. First record for Georgia. Distribution. An extremely rare species known from Bulgaria, Georgia, Greece, Macedonia, Serbia, and Romania (Wagner and Joost 1988;Oboňa et al. 2019a;Ježek et al. 2020 Distribution. 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 (Ježek 2006;Omelková and Ježek 2012;Ježek et al. 2013Ježek et al. , 2018b.
Note. First record for Azerbaijan and Georgia.

Pneumia gracilis gracilis (Eaton, 1893)
Material examined. Georgia: G 02, 24.9.2019, 1♂, slide Inv. No. 25585, leg. TK, PM. Distribution. 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 (Ježek 2002(Ježek , 2004Ježek et al. , 2017Ježek and Omelková 2012;Omelková and Ježek 2012  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.
Note. First record for Azerbaijan.
Note. First record for Azerbaijan. First record for Georgia, from a territory other than Abkhazia.  The number of sites where the species is present is given in brackets after the species name. When species cooccurred at the locality, an offset is also shown in the legend and described in the Materials and methods. 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. Distribution. 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 (Ježek and Goutner 1995;Krek 1999;Ježek 2002;Kvifte et al. 2011Kvifte et al. , 2013Ježek and Omelková 2012;Salmela et al. 2014;Ježek et al. 2017;Oboňa et al. 2019a).
Note. First record for Azerbaijan. It is a vector of larval stages of Rhabditis nematodes and Gamasidae mites .

Saraiella rotunda
Note. First record for Azerbaijan, Georgia, and Transcaucasia.
Note. 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. Distribution. Species known only from the original paper from Armenia .

Threticus balkaneoalpinus
Note. First record for Georgia, territory other than Abkhazia.
Note. 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.
Note. 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.
Note. 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. Distribution. Species known only from the original paper from Abkhazia (Ježek 1987; only on the basis of a holotype (male) from Kolchidian lowland (Novyj Afon) at the Black Sea coast).

Yomormia afonensis
Note. 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.).

Discussion
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 (Ježek et al. , 2018Oboňa et al. 2017Oboňa et al. , 2019a, each of our subsequent expeditions has highlighted species new for science and a number of new records for the Caucasus (or Transcaucasia) or for countries where we carried out the sampling. This study is no exception, and in addition to species new to science (Ježek et al. 2021), we provide faunistic data on 46 species of the family Psychodidae in this publication, up to 12 of which are first records for Transcaucasia, 22 for Azerbaijan, and 17 for Georgia. A total of 80 sampling sites was sampled during the 2019 campaign (24 in Azerbaijan and 56 in Georgia). Therefore, this is another significant shift in the knowledge of the faunistics of this family, but also in regard to the biogeographical and ecological aspects of several recorded species.
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 (Ježek 1988), of which we recorded only other one male. Other extremely rare species known only from Abkhazia until now are Threticus petrosus Ježek, 1997, now known from only two sites, and Yomormia afonensis Ježek, 1987, which has been found at only three localities. The species Tonnoiriella arcuata Ježek, 1997, was also very rare in Abkhazia, but we found this species at five other sites located at a large range of elevations in the Greater Caucasus. Other species were not considered rare in Abkhazia, for example Sycorax caucasica Ježek, 1990, Mormia ckvitariorum Ježek, 1987, and Seoda svanetica (Ježek, 1988), but we found them at only one site and only one (S. caucasica, S. svanetica) to three (M. ckvitariorum) males. This could be caused by the great rarity of these species in other parts of Transcaucasia or in the change that the fauna has undergone under the influence of various factors during the 30 years that divide the records from Abkhazia and our recent ones. However, these assumptions are based on the results obtained using the collection methods described above. Therefore, in order to answer this question definitively, it is necessary to carry out research using more appropriate collection methods and corresponding statistical analyses.
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. 8) and G 09 (Fig. 9), which are located at relatively low elevations along watercourses flowing through preserved deciduous forests (G 09 in Nature Reserve) and characterized by a high diversity of habitats. Also, most of the other sampling sites with high diversity are located in forests (e.g., A 02, G 09, G 40) or rich lush shrub vegetation (e.g., A 05). It is also interesting that many of the first records and records of rare species were obtained from the above-mentioned (close to) natural sites with highest species richness (A 02 with seven spp, and A 23 and G 09 with nine spp each). 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.