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

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. Jan Ježek, Peter Manko, Jozef Oboňa.

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 long‐term 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 . For the above reasons, this is another contribution to knowledge of the : , especially , and from Azerbaijan and Georgia.

is a relatively species-rich family, with nearly 500 species known to occur in Europe and the adjacent island areas; for more details, see Wagner (2018). The following authors contributed considerably to knowledge on the Transcaucasian : Perfiľev (1966); Lewis (1982); Artemiev and Neronov (1984); Vaillant (1972–1983); Vaillant and Joost (1983); Wagner (1981, 1990, 2018); Wagner and Joost (1986, 1988); Gugushvili and Lomtadze (2002); Wagner et al. (2002); Ježek (1987, 2004, 2006a, 2007, 2009); Melaun et al. (2014); Ježek et al. (2014, 2018) and recently Oboňa et al. (2017, 2019a).

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 (Hrivniak et al. 2020). The parts of Georgia and Azerbaijan sampled in this research cover parts of the Greater Caucasus and Lesser Caucasus (see Fig. 1), separated by the Rieni and Kura river valleys and partly connected in the Likhi (Surami) mountain range at a relatively high elevation, with the lowest point of 949 m a.s.l. (Surami Pass).

Figure 1.

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 (Ježek et al. 2018, 2021; Oboňa et al. 2017, 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 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 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 (Inv. No.) and are included in the collection of National Museum Prague collections (), see Tkoč et al. (2014). The nomenclature is modified from Vaillant (1972–1983) and Wagner (1990, 2018) using the classifications of Ježek (1990a, 2007); Ježek and van Harten (2005); Omelkova and Ježek (2012); Oboňa and Ježek (2014) and Kroča and Ježek (2015, 2019). We use the term “Transcaucasia” (Armenia, Azerbaijan, and Georgia) according to the Catalogue of Palaearctic ; 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:

Tibor Kovács;

Dávid Murányi;

Gilles Vinçon.

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.

Figure 2.

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.

Figure 7.

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.

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.

Family

Subfamily

Ježek, 1990

7AA20BE6-35DF-59B2-AF12-BD3FF0DC5C9C

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).

Haliday in Curtis, 1839

C1A8651D-80BF-55CA-BA3D-AAF352278776

Azerbaijan: A 02, 9.5.2019, 1♂, slide Inv. No. 25663, leg. JO (Fig. 2).

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).

First record for Azerbaijan and Transcaucasia.

Wagner, 1981

9B97C88B-D721-5A32-B30A-C402B053449B

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. 2).

Rare Caucasian species known only from a few localities (Wagner 1981; Oboňa et al. 2019).

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.

(Tonnoir, 1940)

DA18CB24-53D4-50A0-87AB-63238E0659C6

Georgia: G 03, 27.9.2019, 1♀, slide Inv. No. 25604, leg. PM (Fig. 2).

Coprophagous species with a Western Palaearctic distribution. Known from Europe (Belgium, Great Britain, France, Germany, Ireland, the Netherlands, and Norway) (Ježek et al. 2018b; Wagner 2018).

First record for Georgia and Transcaucasia.

(Tonnoir, 1919)

15A297F7-E9A2-5D54-85A7-4AA0017F5F02

Georgia: G 09, 2.5.2019, 1♀, slide Inv. No. 25731, leg JO (Fig. 2).

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 (Ježek and Omelková 2012; Oboňa et al. 2019b).

First record for Georgia and Transcaucasia.

(Tonnoir, 1940)

630802C1-AFD3-5D09-85D4-8E4C3F4F694D

Georgia: G 28, 6.7.2019, 1♂, slide Inv. No. 25609, leg. PM (Fig. 2).

Generally common species, occurring across Europe to Transcaucasia, associated with fungi (Kroča and Ježek 2019).

First record for Georgia, from a territory other than Abkhazia.

(Zetterstedt, 1850)

8855005F-707E-5CEF-88B8-0644C89DAB75

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. 2).

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 (Ježek and Yağci 2005; Kvifte 2012; Afzan and Belqat 2016; Ježek et al. 2018).

First record for Georgia.

(Quate, 1955)

C444E378-73CF-593F-B05D-53A81ABC431E

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. 2).

Common Holarctic species, eurybiotic (Kroča and Ježek 2019).

(Kincaid, 1899)

3850957D-26CC-515C-8B9F-8E04D7F6B511

Azerbaijan: A 22, 5.5.2019, 1♀, slide Inv. No. 25626, leg. JO (Fig. 2).

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 (Andersen and Håland 1995; Ježek 2003; Kvifte et al. 2011; Kvifte and Andersen 2012; Elgueta and Ježek 2014; Haselboeck 2016; Ježek et al. 2018b; Oboňa and Kozánek 2018; Wagner 2018).

First record for Azerbaijan and Transcaucasia.

Ježek, 1983

F337EEE0-C9DA-5544-9894-FE37F64B4FCA

Georgia: G 23, 29.4.2019, 1♀, slide Inv. No. 25714, leg. JO (Fig. 2).

Common European and Western Siberian species recorded from Belgium, Czech Republic, Greit Britain, Slovakia, Slovenia, and the Netherlends (Omelková and Ježek 2012; Ježek et al. 2018b; Wagner 2018).

First record for Georgia and Transcaucasia.

Ježek, 1987

40372BF0-BB2A-512C-988B-FEC58A759490

Azerbaijan: A 14, 7.5.2019, 3♂♂, slide Inv. No. 25738, 25739 and 25740, leg. JO (Fig. 3).

Figure 3.

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. The number of sites where the species is present is given in brackets.

Species known only from the original description from Abkhazia (Ježek 1987). First record since the original description, first record for Azerbaijan. Species known only from lowland and foothills of the Greater Caucasus.

(Ježek, 1983)

77159C59-3563-5287-BE24-A1C5DCBF8AB1

Azerbaijan: A 02, 9.5.2019, 1♂, slide Inv. No. 25661, leg. JO (Fig. 3).

Species known only from Czech Republic, Greece, Slovakia, and Slovenia; until now considered as a rare European species (Ježek 1983; Omelková and Ježek 2012; Oboňa and Ježek 2014).

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.

Krek, 1971

19F78687-6D71-5F77-9269-E82EE27E749A

Azerbaijan: A 23, 6.5.2019, 1♂, slide Inv. No. 25693, leg. JO (Fig. 3). Georgia: G 12, 9.7.2019, 1♂, slide Inv. No. 25613, leg. PM, GV; G 32, 2.7.2019, 1♂, slide Inv. No. 25615, leg. TK, DM, GV; G 40, 13.7.2019, 1♂, slide Inv. No. 25634, leg. TK, PM, DM, GV.

Locally common species, known from several European countries: Belgium, Austria, Great Britain, Ireland, Bosnia and Herzegovina, Czech Republic, Greece, and Macedonia (Omelková and Ježek 2012; Ježek et al. 2018b).

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.

(Walker, 1856)

45B28BBF-0AE6-5334-9620-4CC8CDB87B4D

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. 3).

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 (Ježek and Yağci 2005; Salmela et al. 2014; Kvifte et al. 2016; Ježek et al. 2018a; Oboňa et al. 2019a).

First record for Georgia.

Ježek, 1990

8C1AF31E-2EBE-5644-A711-C9D49CE4B58F

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. 3).

Vaillant, 1983, was described from Schelda nr. Kashadl (Central Caucasus: Georgia, probably Gora Shkhel’da, omitted (overlooked) from Georgia in Oboňa et al. 2019a); the subspecies is known only from the original description (Ježek 1990b) from S. E. Iran, Kerman province, Kuh-e Lalehzar (top m a.s.l. 4374, 3850–4374 m), holotype and 9 paratypes (males).

Vaillant, 1983 was raised to genus level by Ježek (2001) after previously being considered a subgenus of Vaillant, 1961. This is the first record since the original description.

Eaton, 1893

87AA84FF-7E35-572E-B569-1909E76C8C4B

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 (Kroča and Ježek 2019).

Eaton, 1893

F0520A8E-B010-50D2-BF67-C42C139FF9AA

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 (Kroča and Ježek 2019).

First record for Georgia, from a territory other than Abkhazia.

Kvifte, 2010

2991DEDC-BB5F-5286-8608-8836F1FD74D2

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 (Ježek 2009; Kvifte 2010; Kvifte et al. 2011; Ježek et al. 2017).

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.

Krek, 1966

18CAD882-D14F-5203-B938-E470F7F638AC

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 (Krek 1966, 1967a, 1967b, 1970, 1972, 1974, 1979a, 1999; Krek et al. 1976; Mučibabič et al. 1984; Vaillant 1978; Wagner and Joost 1988; Wagner 1990; Ježek et al. 2018; Oboňa et al. 2019a).

First record for Georgia.

Eaton, 1893

FB7404DD-E6AA-5EDB-A5CE-74DA43B7F99A

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 (Ježek 2004, 2009; Kvifte et al. 2013; Wagner 2013; Oboňa and Ježek 2014; Ježek et al. 2017; Oboňa et al. 2019a).

First record for Georgia.

Vaillant, 1978

A85D87E2-E7FA-5428-B9F6-162AA66DAC46

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.

Vaillant, 1978

617E1834-269C-5EE8-A848-A6E3985BB567

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 (Wagner and Joost 1988; Oboňa et al. 2019a; Ježek et al. 2020).

First record for Azerbaijan.

Tonnoir, 1940

8B576D47-BD3D-5DF5-A4AA-4F790DA35ED4

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 (Ježek et al. 2018b; Wagner 2018).

First record for Georgia and Transcaucasia.

(Meigen, 1804)

DB7C6AF7-08A0-5BAF-8E90-87C62061F86C

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 (Ježek 2006; Omelková and Ježek 2012; Ježek et al. 2013, 2018b).

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.

(Eaton, 1893)

0A5DC85E-BD94-5355-80BB-99E617E6182C

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 (Ježek 2002, 2004; Ježek et al. 2012, 2017; Ježek and Omelková 2012; Omelková and Ježek 2012).

First records for Georgia outside of Abkhazia.

(Meigen, 1818)

D4D81DFF-CF9F-5D43-8F82-25FB89D88D49

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 (Wagner 1981; Ježek and Goutner 1995; Krek 1999; Ježek 2002; Omelková and Ježek 2012; Kvifte et al. 2013; Ježek et al. 2017, 2018a).

First record for Azerbaijan.

(Meigen, 1804)

6A9D0B55-3FB6-5196-9DCF-FE6C2B0371FB

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 (Kroča and Ježek 2019; Oboňa et al. 2019).

(Tonnoir, 1940)

F3427769-B8AB-511D-AA57-0B4121DB1E04

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 (Kroča and Ježek 2019; Oboňa et al. 2019a).

(Eaton, 1893)

8F7362A2-E887-529D-AA04-5FDA8FEE0AF1

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 (Krek 1999; Ježek 2002; Kvifte et al. 2011, 2013; Omelková and Ježek 2012; Ježek et al. 2017).

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.

(Tonnoir, 1922)

68F71813-E169-5627-835F-D0888D9DC3CF

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 (Kroča and Ježek 2019; Oboňa et al. 2019a).

(Linné, 1758)

BB20353A-CC95-5088-91E3-643F75B8A914

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., Ježek 1990b; Oboňa and Ježek 2014; Oboňa et al. 2019a).

First record for Azerbaijan.

(Banks, 1894)

33C1F923-768D-5F1B-9124-D53D6F5F8279

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 (Krek 1985; Wagner 1990, 2018; Ježek and Goutner 1995; Ježek and Yağci 2005; Kvifte et al. 2011; Salmela et al. 2014; Ježek et al. 2017).

First record for Azerbaijan. First record for Georgia, from a territory other than Abkhazia.

(Eaton, 1904)

3828E873-A56E-5E71-9F31-88E14080E682

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 (Ježek and Goutner 1995; Krek 1999; Ježek 2002; Kvifte et al. 2011, 2013; Ježek and Omelková 2012; Salmela et al. 2014; Ježek et al. 2017; Oboňa et al. 2019a).

First record for Azerbaijan.

(Vaillant, 1988)

33904ADD-0DEA-5925-8977-8DC5099E364A

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 (Ježek and Omelková 2012; Ježek et al. 2017).

First record for Azerbaijan, Georgia, and Transcaucasia.

(Tonnoir, 1922)

7CEA480C-520B-5F43-81B1-E99A4354A96D

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 (Ježek 1990c; Wagner 2013; Salmela et al. 2014; Ježek et al. 2017; Oboňa et al. 2019a).

First record for Azerbaijan. It is a vector of larval stages of nematodes and mites (Ježek and Omelková 2012).

Wagner, 1981

EC8AE2A0-B482-5390-BE5F-EA33C55F519B

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; Ježek et al. 2018). First record for Georgia.

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.). 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 Ježek, 1990 is thus sunk as a synonym herein.

(Krek, 1970)

770EAF0E-3A29-5B60-8519-0C26F5AA7306

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 (Ježek 2006; Oboňa et al. 2019b). Occurs in spring areas and swamps, forest edge in Slovakia (High Tatras Mts.) (Ježek 2006).

First record for Azerbaijan, Georgia, and Transcaucasia.

(Ježek, 1988)

785DAD40-2B24-5A41-A534-F61C157772D5

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. (Ježek 1988 as ).

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.

Oboňa & Ježek, 2017

7D47C182-E67C-5C59-A1A1-1160B31E8145

Azerbaijan: A 05, 1.10.2019, 1♂, slide Inv. No. 25575, leg. PM.

Species known only from the original paper from Armenia (Oboňa et al. 2017).

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.

Krek, 1972

0782973D-0244-5717-A669-71A51EF760AA

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 (Krek 1990; Ježek 1995; Withers 1997; Ježek and Omelková 2007; Oboňa and Ježek 2014; Oboňa et al. 2019b).

First record for Georgia, territory other than Abkhazia.

Ježek, 1997

0BDD7D38-2FED-521A-BDFC-2A498D4DC2F3

Georgia: G 06, 24.9.2019, 1♂, slide Inv. No. 25598, leg. GV.

Known only from the original paper from Abkhazia (Ježek 1997; Bzybskij khrebet, pastoral community Kot-Kot nr. peak Khimsa (3033 m a.s.l.) – holotype + 9 paratypes (males)).

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.

(Say, 1824)

38FFC610-53BF-50CE-AC93-10E49FFAD9C4

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 (Kroča and Ježek 2019).

Ježek, 1997

EDBEBC77-50D2-5348-8DB0-EF614CCF4142

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 (Ježek 1997; Pskhu), holotype + 6 paratypes (males).

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.

(Eaton, 1893)

18282418-0BAA-5FB0-A92A-66697C738834

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 (Ježek and Omelková 2012; Salmela et al. 2014; Ježek et al. 2017, 2018a).

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.

Ježek, 1987

0FE423EB-3D6D-5D1A-A3B5-ACB97D945453

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 (Ježek 1987; only on the basis of a holotype (male) from Kolchidian lowland (Novyj Afon) at the Black Sea coast).

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 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. 2014, 2018; Oboňa et al. 2017, 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 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 (Ježek 1988), of which we recorded only other one male. Other extremely rare species known only from Abkhazia until now are Ježek, 1997, now known from only two sites, and Ježek, 1987, which has been found at only three localities. The species 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 Ježek, 1990, Ježek, 1987, and (Ježek, 1988), but we found them at only one site and only one (, ) to three () 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: and occur at large range of elevations, and prefer low elevations in lowlands and low foothills, elevations higher than 1000 m, and 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 (, , ).

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).

Figure 8.

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.

Figure 9.

The collecting site with the highest species richness in Georgia (G 09), Batsara River and its side brook, Kakheti region, Batsara Nature Reserve, with nine species collected; photograph P. Manko.

Conclusions

Out of the total number of 46 species/subspecies ( 1 sp., 1 sp. 44 sp.) and 182 slides, 12 species are recorded for the first time for Transcaucasia (namely , , , , , , , , , , , ), 22 species for Azerbaijan (namely , , , , , (.) , , , , , , , , , , , , , , , , ), 17 species for Georgia (, , , , , , , , , , , , , , , , ), and nine species for the first time since their original descriptions (, , , , , , , , ). 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.