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Research Article
A contribution towards checklist of fungus gnats (Diptera, Diadocidiidae, Ditomyiidae, Bolitophilidae, Keroplatidae, Mycetophilidae) in Georgia, Transcaucasia
expand article infoOlavi Kurina‡
‡ Estonian University of Life Sciences, Tartu, Estonia

Corresponding author: Olavi Kurina ( olavi.kurina@emu.ee )

Academic editor: Vladimir Blagoderov
© 2021 Olavi Kurina.
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: Kurina O (2021) A contribution towards checklist of fungus gnats (Diptera, Diadocidiidae, Ditomyiidae, Bolitophilidae, Keroplatidae, Mycetophilidae) in Georgia, Transcaucasia. ZooKeys 1026: 69-142. https://doi.org/10.3897/zookeys.1026.63749
ZooBank: urn:lsid:zoobank.org:pub:05EFF10E-6214-4368-BE47-1AA57A2C38D7
Open Access

Abstract

The fungus gnats of Georgia are studied based on 2682 specimens collected from 57 localities during 2011–2019. Altogether, 245 species are recorded including four species of Bolitophilidae, three species of Diadocidiidae, two species of Ditomyiidae, 34 species of Keroplatidae and 202 species of Mycetophilidae. 230 and 188 species are recorded from Georgia and the whole of Transcaucasia for the first time, respectively. Three new species – Sciophila georgei sp. nov., Leia katae sp. nov. and Anatella metae sp. nov. – are described including detailed illustrations of the male terminalia. Photographs are provided for an additional 38 species to highlight a variability of their general facies. Combined with earlier published data, the number of fungus gnat species in Georgia is set at 246. The estimated diversity of fungus gnats in Georgia is calculated using non-parametric methods and discussed with respect to other Western Palaearctic regions.

Keywords

Fungus gnats, Georgia, new species, Sciaroidea, species diversity, taxonomy, Transcaucasia

Introduction

The last decades can be characterized by an upturn of systematics, taxonomy and biodiversity studies (e.g. Padial et al. 2010; Kõljalg et al. 2020; Wheeler 2020). That is also true in the case of the insects order Diptera (e.g. Wiegmann et al. 2011; Kirk-Spriggs and Sinclair 2017; Borkent et al. 2018) including the superfamily Sciaroidea (e.g. Kjaerandsen et al. 2007; Borkent and Wheeler 2012; Ševčík et al. 2013; Fitzgerald and Kerr 2014). Seven families and a insertae sedis group are included in Sciaroidea (Ševčík et al. 2016; Mantič et al. 2020), whereas five of them, viz. Diadocidiidae, Ditomyiidae, Bolitophilidae, Keroplatidae and Mycetophilidae are conjoined under a common name ‘fungus gnats’. Today, more than 5,500 species of fungus gnats are known globally (Evenhuis and Pape 2021; Fungus Gnats Online Authors 2021), however, their actual diversity is insufficiently known, especially in tropical regions of the world. As expected, the group is best studied in Europe with about 1,200 named species (Chandler 2013) yielded by more than 200 years of studies pioneered by the “father” of dipterology J.W. Meigen (e.g. Meigen 1804, 1818). Nevertheless, even in Europe, new species are described annually and e.g. in Nordic countries nearly 120 new species are waiting to be described (Kjærandsen and Søli 2020). While fungus gnats are mostly forest dwellers preferring shady and humid habitats, some species are also recorded from more open landscapes (Falk and Chandler 2005). They are small to medium size nematocerous flies with a humpbacked habitus, prominent coxae and hyaline or patterned wings (see e.g. Figs 8, 9). The trophic strategy of fungus gnats is diverse: the majority of the known associations are those with fungal fruiting bodies or mycelium-penetrated forest litter including decaying wood but several species develop in other terrestrial habitats and/or can also be sporophagous or predators in the larval stage (e.g. Matile 1997; Ševčík 2010; Jakovlev 2012; Põldmaa et al. 2016; Mantič et al. 2020).

Transcaucasia, the area southwards from the Greater Caucasus Mountains that includes the countries of Georgia, Azerbaijan and Armenia, is considered one of the biodiversity hotspots of the world, with a remarkable number of endemic species (Myers et al. 2000). However, limited attention has been paid to the biodiversity research in the area so far (Mumladze et al. 2020) and most organism groups, including Diptera and fungus gnats in particular, are rather superficially studied. There are 33 species of fungus gnats recorded from Azerbaijan (Zaitzev 1994, 2003; Zaitzev and Ševčík 2003) and seven species from Armenia (Joost and Plassmann 1985, Zaitzev 1994). From Georgia, only one species was known (Zaitzev 1994) prior to Kurina and Jürgenstein (2013) who described two new Orfelia Costa (Keroplatidae) species from Marelisi, NW of Borjomi. Later on, Jürgenstein et al. (2015), Kurina et al. (2015), Kurina (2018), Thormann et al. (2019) and Ševčík et al. (2020) provided data on another twelve species and the number of fungus gnat species from Georgia is currently set at 15. Furthermore, an additional 24 fungus gnat species have been listed to occur in Transcaucasia but without a specified region (Zaitzev 1994, 2003). Concerning neighbouring areas, 91 species of fungus gnats are recorded from the northern slopes of the Great Caucasus ridge, most of them from the surroundings of Mt Elbrus (Joost and Plassmann 1976, 1979, 1985, 1992, Plassmann 1976).

During the last decade, a considerable amount of fungus gnat material from Georgia has accumulated in the author’s possession. The aim of the current contribution is to provide results of the study based on that material along with summarising all available published information on Georgian fungus gnats.

Material and methods

The material was collected from 2011 to 2019 using different methods in the course of 61 collecting events from 57 localities in Georgia (Table 1, Fig. 1). The majority of the material was collected sweeping during three expeditions by the author in May of 2012 and 2013 and August-September 2014. Additional material from Malaise trap samples is included from the provinces Samegrelo-Zemo-Svanethi, Imereti and Kaheti; a sporadic material as a by-product of light trap collecting is also included (Table 1). The collecting localities (see Fig. 2 for examples) varied from more open landscape in Vardzia (Table 1: SJ-10) to highly forested mountain areas in Kintrishi (Table 1: A-5–8), Mtirala (Table 1: A-1–4) and Borjomi-Kharagauli (Table 1: I-5–17) National Parks, and subalpine areas in the surroundings of Stepantsminda (Table 1: MM-1–6), Bakuriani (Table 1: SJ-6–7) and Ushguli (Table 1: SZS-2–3).

Table 1.
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Collecting data of fungus gnats in Georgia arranged by administrative regions of the country. Codes for the collecting events are used on the Figure 1 and within the list of species. The asterisk (*) indicates collecting with a Malaise trap for which the exact collecting dates are provided in the text.

Region Locality Coordinates Altitute (m) Collecting date(s) Collecting method Collector Code
Samegrelo-Zemo Svanethi Mestia 43°02.97'N, 42°44.72'E 1500 28.vii.2017 light trap O. Kurina SZS-1
Chvabiani 43°02.47'N, 42°51.03'E 1630 29.vii.2017 light trap O. Kurina SZS-2
S of Lakhushdi, meadow 42°59.93'N, 42°39.02'E 1270 13–14.vi.2019 Malaise trap X. Mengual SZS-3
near Ushguli, path to glacier 42°56.62'N, 43°03.23'E 2220 15–17.vi.2019 Malaise trap X. Mengual SZS-4
Adjara Mtirala NP, visitor centre 41°40.65'N, 41°51.30'E 240 19.v.2013 sweeping O. Kurina A-1
Mtirala NP, visitor centre 41°40.65'N, 41°51.33'E 230 19.v.2013 at light O. Kurina A-2
Mtirala NP, visitor centre 41°40.35'N, 41°52.53'E 270 20.v.2013 sweeping O. Kurina A-3
Mtirala NP, visitor centre 41°40.91'N, 41°50.70'E 220 20.v.2013 at light O. Kurina A-4
Kintrishi NP 41°45.76'N, 41°58.67'E 320 21.v.2013 sweeping O. Kurina A-5
Kintrishi NP 41°45.76'N, 41°58.67'E 320 21.v.2013 at light O. Kurina A-6
Kintrishi NP 41°45.20'N, 41°58.63'E 450 22.v.2013 sweeping O. Kurina A-7
Kintrishi NP 41°46.40'N, 41°58.08'E 460 22.v.2013 sweeping O. Kurina A-8
Imereti Chiatura 42°17.00'N, 43°17.00'E 480 17.v.2011 light trap U. Jürivete I-1
Tshunkuri 42°24.00'N, 42°35.00'E 230 20.v.2011 light trap U. Jürivete I-2
Patara Vardzia, W of Kharagauli 42°0.55'N, 43°04.62'E 740 v-x.2013* Malaise trap O. Kurina I-3
Patara Vardzia, W of Kharagauli 42°01.32'N, 43°11.10'E 370 18.v.2013 sweeping O. Kurina I-4
Marelisi 41°57.07'N, 43°17.02'E 430 18.v.2012 sweeping O. Kurina I-5
Marelisi 41°57.93'N, 43°17.35'E 410 19.v.2012 sweeping O. Kurina I-6
Marelisi, on road to railway station 41°58.02'N, 43°17.35'E 440 19.v.2012 at light O. Kurina I-7
Marelisi 41°57.00'N, 43°17.00'E 460 20.v.2012 indoors O. Kurina I-8
Marelisi, on path to hill 41°56.38'N, 43°16.62'E 690 20.v.2012 sweeping O. Kurina I-9
Marelisi 41°56.80'N, 43°17.05'E 450 20.v.2012 sweeping O. Kurina I-10
Marelisi 41°58.02'N, 43°17.38'E 400 17.v.2013 sweeping O. Kurina I-11
Marelisi 41°56.28'N, 43°16.98'E 460 17.v.2013 sweeping O. Kurina I-12
Marelisi 42°56.46'N, 43°17.05'E 460 18.v.2013 al light O. Kurina I-13
Marelisi 41°56.28'N, 43°16.98'E 460 29.viii.2014 sweeping O. Kurina I-14
Marelisi 41°56.38'N, 43°16.47'E 760 30.viii.2014 sweeping O. Kurina I-15
Marelisi, close to railway station 41°58.23'N, 43°18.65'E 400 20.v.2012 al light O. Kurina I-16
Marelisi, close to railway station 41°58.14'N, 43°18.63'E 410 23.v.2013 sweeping O. Kurina I-17
Shida-Kartli W of Surami 42°01.57'N, 43°29.88'E 940 18.v.2012 sweeping O. Kurina SK-1
Samtskhe-Javakheti road from Abastumani to Saime, near river 41°46.63'N, 42°50.23'E 1370 10–11.vi.2019 Malaise trap X. Mengual SJ-1
road from Abastumani to Saime 41°47.82'N, 42°50.63'E 1730 10–11.vi.2019 Malaise trap X. Mengual SJ-2
Borjomi 3 km--W, Likani 41°50.15'N, 43°19.95'E 940 21.v.2012 sweeping O. Kurina SJ-3
Borjomi 3.5 km--W, Likani 41°50.11'N, 43°19.92'E 950 31.viii.2014 sweeping O. Kurina SJ-4
Bakuriani 2 km--NW 41°45.77'N, 43°30.28'E 1630 31.viii.2014 sweeping O. Kurina SJ-5
Bakuriani 5 km--S, road from Bakuriani to Tabatskuri 41°42.33'N, 43°30.13'E 2120 1.ix.2014 sweeping O. Kurina SJ-6
Bakuriani 3 km--SW, road from Bakuriani to Tabatskuri 41°43.33'N, 43°29.87'E 1870 1.ix.2014 sweeping O. Kurina SJ-7
Bakuriani 1 km--SW, road from Bakuriani to Tabatskuri 41°44.22'N, 43°30.75'E 1740 1.ix.2014 sweeping O. Kurina SJ-8
Bakuriani 2 km--NW 41°45.77'N, 43°30.28'E 1630 1.ix.2014 sweeping O. Kurina SJ-9
Vardzia, near Tirebi guesthouse 41°24.17'N, 43°19.23'E 1260 22.v.2012 at light O. Kurina SJ-10
Mtskhetha-Mthianethi Stephantsminda 42°39.28'N, 44°39.28'E 1870 15.v.2012 at light O. Kurina MM-1
Stephantsminda, road to Gegriti Trinity Church 42°39.77'N, 44°37.50'E 1980 16.v.2012 sweeping O. Kurina MM-2
Gvelethi NW of Stepantsminda 42°42.28'N, 44°37.27'E 1640 16.v.2012 at light O. Kurina MM-3
Gvelethi NW of Stepantsminda, surroundings of lake 42°43.37'N, 44°37.12'E 1520 17.v.2012 sweeping O. Kurina MM-4
Gvelethi NW of Stepantsminda, surroundings of waterfall 42°42.23'N, 44°37.20'E 1570 17.v.2012 sweeping O. Kurina MM-5
Stephantsminda, road to Gegriti Trinity Church 42°40.02'N, 44°37.15'E 2090 17.v.2012 at light O. Kurina MM-6
Gudauri 42°26.23'N, 44°29.95'E 1780 8.vii.2019 light trap A. Selin MM-7
Dgnali 42°13.43'N, 44°40.02'E 910 15.v.2012 sweeping O. Kurina MM-8
Mtskhetha-Mthianethi Zaridzeebi 42°42.08'N, 44°54.00'E 870 22.v.2011 light trap U. Jürivete MM-9
Saguramo 41°54.00'N, 44°46.00'E 600 16.v.2011 light trap U. Jürivete MM-10
Saguramo 41°53.07'N, 44°46.78'E 920 15.v.2012 sweeping O. Kurina MM-11
Saguramo 41°53.07'N, 44°46.78'E 920 15.v.2013 sweeping O. Kurina MM-12
Saguramo 41°53.07'N, 44°46.78'E 920 28.viii.2014 sweeping O. Kurina MM-13
Saguramo 41°53.13'N, 44°46.73'E 890 4.ix.2014 sweeping O. Kurina MM-14
Kvemo Kartli Manglisi 6 km–S 41°39.89'N, 44°23.10'E 1190 23.v.2012 sweeping O. Kurina KK-1
Kakheti Dzveli Shuamta W of Telavi 41°54.60'N, 45°24.33'E 1000 2.ix.2014 sweeping O. Kurina K-1
Gurgeniani, W of Lagotekhi 41°52.67'N, 46°14.55'E 630 3.ix.2014 sweeping O. Kurina K-2
Matsimi near Lagotekhi 41°48.55'N, 46°18.73'E 440 3.ix.2014 at light O. Kurina K-3
Lagodekhi NR, near administration building 41°50.50'N, 46°16.98'E 560 28.v–9.vi.2011 Malaise trap G. Japoshvili K-4
Lagodekhi NR, Matsimi river gorge 41°47.75'N, 46°17.12'E 350 17–27.v.2011 Malaise trap G. Japoshvili K-5
Lagodekhi NR (Malaise trap #3) unavailable 15–25.vi.2014 Malaise trap G. Japoshvili K-6
Figure 1. 

Collecting localities in Georgia. The codes are those used in Table 1.

The vast majority of the material was initially preserved in 70% ethyl alcohol where most of it is also stored after determination. Every species per locality is arranged in a separate glass vial equipped with collecting and determination labels. Some specimens were double pinned directly after collecting, whereas part of the initially alcohol-preserved specimens were mounted using the method described by Vockeroth (1966) and double pinned thereafter. The majority of the material was determined directly in alcohol as that also allowed observation of the terminalia. However, in a number of cases a more detailed study of male terminalia proved to be unavoidable. For that, terminalia were detached and treated with about 10% warm potassium hydroxide followed by neutralization with acetic acid and washing with distilled water. Terminalia were studied in glycerine and stored as glycerine preparations in small plastic vials attached to the rest of the specimen (see also Kurina 2008a).

Figure 2. 

A gallery of collecting localities in Georgia. For codes see Table 1 A Mestia (SZS-1) B Mtirala NP (A-1) C Kintrishi NP (A-5) D Marelisi (I-12) E Bakurjani (SJ-6) F Lagotekhi (K-3) G Vardzia (SJ-10) H Gvelethi near Stepantsminda (MM-5) I Dgnali (MM-8).

Illustrations of the terminalia were prepared using a U-DA drawing tube attached to a compound microscope Olympus CX31. The digital images of the general habitus and terminalia were combined using the software LAS V.4.1.0. from multiple gradually focused images taken by a Leica DFC 450 camera attached to a Leica 205C stereomicroscope (see also Jürgenstein et al. 2015). Adobe Photoshop CS5 was used for editing the figures and compiling the plates. The morphological terminology follows Søli (1997, 2017) and that of the male terminalia is explained in Figs 4–7. The estimated species richness according to different non-parametric methods (Fig. 14) is calculated using the software EstimateS, Version 9.1.0. (Colwell 2013).

The material is deposited in the following collections:

IUTG Ilia State University, Tbilisi, Georgia;

IZBE Institute of Agricultural and Environmental Sciences, Estonian University of Life Sciences (former Institute of Zoology and Botany), Tartu, Estonia;

ZFMK Zoological Research Museum Alexander Koenig, Bonn, Germany.

The majority of the studied specimens are deposited in IZBE which is not repeated in the species list for every specimen. However, the material collected by X. Mengual (Bonn, Germany) in 2019 is divided between three institutions and the depository is specified in listed material sections below.

Results

Altogether, 2682 studied specimens were identified to 245 different species, viz. four species of Bolitophilidae, three species of Diadocidiidae, two species of Ditomyiidae, 34 species of Keroplatidae and 202 species of Mycetophilidae including three species described as new to science. One additional species of Keroplatidae was included from the literature data (Zaitzev 1994). Moreover, six additional putatively new Mycetophilidae species were recorded, all represented by singletons, some of them of poor quality. Description of these species is deferred pending additional material to be collected. These six species are not included in the species list but are considered in the species richness calculations and distribution analysis (see Discussion). 230 and 188 species are recorded from Georgia and the whole Transcaucasia for the first time, respectively. In the species list, all available literature sources are cited for the species recorded earlier in Georgia and/or in Transcaucasia generally. The studied material is listed, using abbreviations of collecting events provided in Table 1, followed by total number of studied specimens. Distribution in Georgia is given by administrative provinces and the general distribution by zoogeographical regions or subregions. The latter is provided according to Chandler (2013) and subsequent published information available. Some remarks on distribution and/or taxonomy are included for species of special interest. To illustrate the diverse habitus of recorded fungus gnat species a gallery of photographs is provided (Figs 8, 9, 11, 12). In the list of species, the classification follows Fungus Gnats Online (http://www.sciaroidea.info/) except in two cases. Firsty, the subfamily Platyurinae of Keroplatidae is used according to Mantič et al. (2020). Secondy, Brachycampta Winnertz, 1863 is reinstated to the generic status from a subgenus of Allodia Winnertz, 1863 in accordance with a thorough study by Magnussen (2020); this opinion is also implemented in the recent checklist of fungus gnats of Norway by Kjærandsen and Søli (2020).

The new species

Sciophila georgei sp. nov.

http://zoobank.org/D9E0ED72-E487-480C-A89F-4E6DDD98C406
Figs 3A, 4A–G

Type material

Holotype. Male, Georgia, Kakheti, Lagotekhi NR, Matsimi river gorge, 41°47.75'N, 46°17.12'E, 350 m a.s.l., 17–27.v.2011, Malaise trap, leg. G. Japoshvili [see Table 1: K-5] (mounted from alcohol, IZBE). Paratype. Male, same as holotype (mounted from alcohol, IZBE).

Diagnosis

Sciophila georgei sp. nov. can be distinguished by combination of the characters of the male terminalia as follows: lateral branch of gonostylus ventrally with two apical spine-like setae, small internal branch of gonostylus with one spine-like seta, tergite 9 large with medially rising apical margin that bears two large and simple setae, parameres straight and long, extending over tergite 9 apically, aedeagus small, star-shaped.

Description

Male. Body length 2.7–2.8 mm (n = 2). Coloration. Head with vertex and frons dark brown, face and clypeus brown and mouthparts including palpus pale yellow. Scape and pedicel yellow. First three or four flagellomeres yellowish, rest of flagellomeres light brown. Scutum entirely dark brown, antepronotum and proepisternum yellowish, anepisternum, anepimeron and katepisternum light brown, laterotergite and mediotergite brown, scutellum brown. Thoracic setae all yellowish. Wing hyaline, all veins brown including radial veins somewhat darker. Halter with stem and knob pale yellow. All coxae, femora and tibiae yellow, tarsi yellow but seem darker because of dense brown setae. Tibial setae brown, spurs yellowish. Abdomen with tergites light brown, 1–3 tergites somewhat lighter, all sternites yellowish. Abdominal vestiture yellow. Terminalia brown. Head. Ocelli in a shallow triangular arrangement. Medial ocellus somewhat smaller than laterals. Frontal furrow complete. Clypeus subrounded, about as long as broad. Fourth flagellar segment about as long as wide, apical flagellar segment 2.25 times as long as wide basally. Flagellar segments with dense yellowish short setae. Thorax. Scutum covered with short setae, with marginal and prescutellar setae stronger. Antepronotum with 8–9 setae. Proepisternum with 6–7 setae. Anepisternum with 5–6 setae on upper part, katepisternum and anepimeron non-setose. Laterotergite with 7–9 setae on posterior half. Mediotergite with 10–15 setae on lower part. Metepisternum with setulae. Scutellum with setulae and marginal setae not arranged in pairs. Wing. Length 2.5–2.8 mm, length to width 2.4–2.7. Wing membrane uniformly covered with micro- and macrothichia. All veins setose, except sc-r, Rs, R2+3. Costa reaches about one fifth from R4+5 to M1. Sc ending on C before level of furcation of posterior fork. Sc-r located slightly before Rs. r-m about two times as long as m-stem. M4 basally very faint or shortly interrupted at base. Legs. Ratio of femur to tibia for fore, mid and hind legs: 0.83–0.93; 0.89–0.97; 0.84–0.92. Ratio of tibia to basitarsus for fore, mid and hind legs: 1.26; 1.42–1.65; 1.33–1.37. Fore tibia with a spur 2.29–2.81 times of tibial maximum width. Mid tibia with anterior spur 3.08–3.15 times and posterior spur 3.42–3.69 times of tibial maximum width. Hind tibia with anterior spur 2.50–2.60 times and posterior spur 3.47–3.57 times of tibial maximum width. Terminalia (Fig. 4A–G). Gonocoxites fused for short distance ventrobasally forming medial triangular lobe with medial more sclerotized longitudinal ridge internally. Ventromedial margin of gonocoxite with a membranous flange drawn medially out into digitate apically hooked lobe. Gonocoxite covered with uniform setae except non-setose lateroapical and dorsomedial marginal areas. Dorsoposterior margin of gonocoxite with two prominent medially directed setae. Gonocoxal apodeme anteriorly enlarged, shoe-shaped, well discernible in dorsal view. Tergite 9 large, slightly convergent posteriorly, extending over gonocoxites, subapically constricted with two prominent simple setae apically, apical margin medially rising. Parameres long and straight, apically slightly widening, extending over tergite 9 apically. Aedeagus small, star-shaped, medially with posteriorly projecting digitate process. Lateral branch of the gonostylus laterally setose with aggregation of spine like setae along posterior margin; ventral part extended with two prominent apical spine-like setae. Medial branch of gonostylus with 25–30 furcated megasetae. Small internal branch of gonostylus with one prominent medially directed seta.

Figure 3. 

Habitus photos of new species A Sciophila georgei sp. nov., paratype B Anatella metae sp. nov., paratype C Leia katae sp. nov., holotype, terminalia detached. Scale bars: 1 mm.

Female. Unknown.

Etymology

The species is named in honour of Prof. George Japoshvili (Tbilisi, Georgia) in recognition of his contribution to study of the insects’ diversity in Georgia and his invaluable help in collecting the fungus gnat material that underlies the current communication. He was also the collector of the type material of this species.

Figure 4. 

Sciophila georgei sp. nov., male terminalia A dorsal view B ventral view C lateral view D apical part of tergite IX, dorsal view E dorsal view, tergite IX removed F aedeagal complex, dorsal view G gonostylus, internal view H gonostylus, posterior view. Abbreviations: aed = aedeagus, cer = cercus, gc = gonocoxite, gc ap = gonocoxal apodeme, gst lbr = lateral branch of gonostylus, gst mbr = medial branch of gonostylus, gst sibr = small internal branch of gonostylus, par = paramere, tg 9 = tergite IX. Scale bars: 0.1 mm.

Remarks

More than 190 species of Sciophila Meigen are known wordwide (Kurina 2020a, Taber 2021); the most comprehensive key to the Holarctic species is still that by Zaitzev (1982). Fortunately, all subsequently described species are supplemented with appropriate illustrations of the male terminalia (e.g. Polevoi 2001; Salmela and Kolcsár 2017; Taber 2021) that provides an adequate compendium of the morphological distinctions. Following the key by Zaitzev (1982), the new species runs to couplet 31 because of (1) wing with both macro- and microtrichia, (2) gonostylus without additional branches, (3) lateral branch of the gonostylus with two large setae ventroapically, (4) small internal branch of the gonostylus with one large seta, and (5) gonocoxites dorsoapically without protruding appendages. However, S. georgei sp. nov. differs from the species included in this couplet by details in the male terminalia. Notably, by the characters of tergite 9 (large, posteriorly convergent, extends over gonocoxites, bears two prominent simple setae apically, apical margin medially rising) and parameres (long, straight, extending over tergite 9 apically).

Leia katae sp. nov.

http://zoobank.org/FDD299DF-4281-4DD0-9BAC-8B6050D98226
Figs 3C, D, 5A–D, 6A–C

Type material

Holotype. Male, Georgia, Shida-Kartli, W of Surami, 42°01.57'N, 43°29.88'E, 940 m a.s.l., 18.v.2013, sweeping, leg. O. Kurina [see Table 1: SK-1] (mounted from alcohol, IZBE). Paratype. Male, GEORGIA, Samegrelo-Zemo-Svanethi, near Ushguli, path to glacier, 42°56.62'N, 43°03.23'E, 2220 m a.s.l., 15–17.vi.2019, Malaise trap, leg. X. Mengual [see Table 1: SZS-4] (in alcohol, ZFMK)

Diagnosis

Leia katae sp. nov. can be distinguished by the combination of characters as follows: thorax bicolored (scutum yellow, with brown longitudinal stripes; katepisternum with lower half brown), wing tinged yellowish, with faint preapical brownish band, male terminalia with bipartite gonostylus (lateral prong shorter, convolute and apically hooked; medial prong longer, tapering with preapical small tooth at ventral margin).

Description

Male. Body length 6.7–6.9 mm (n = 2). Coloration. Head with vertex brown, frons yellow, face, clypeus and mouthparts including palpus pale yellow. Scape and pedicel pale yellow. First two flagellomeres yellowish, flagellomeres 3–14 brown. Thorax bicoloured: scutum yellow with three brown longitudinal stripes, which are posteriorly completely fused, lateral stripes begin at a distance of one third from anterior margin, medial stripe shortly split anteriorly, lateral parts of scutum yellow; antepronotum, proepisternum and anepisternum yellow, posterior margin of anepimeron light brown, katepisternum with lower half brown and upper half yellow, laterotergite brown with posterior half yellowish, mediotergite brown, scutellum basally yellowish, apically brown. Thoracic setae all yellow. Wing with yellowish tinge and preapical very faint transverse brownish band reaching to M2, all veins yellowish including radial veins somewhat darker. Halter with stem and knob pale yellow. All coxae and femora yellow, except hind femur apically with narrow brown band, all tibiae, tarsi yellow but seem darker because of dense brown setae. Tibial setae brown, spurs yellowish. Abdomen with all tergites brown and sternites yellow. Abdominal vestiture yellow. Terminalia brown with gonocoxite medially and gonostylus anteriorly yellow. Head. Ocelli in a linear arrangement. Medial ocellus about twice smaller than laterals, which are separated from eye margins by less than their own diameter. Frontal furrow complete. Clypeus obovoid. Fourth flagellar segment about as long as wide, apical flagellar segment 2.5 times as long as wide basally. Flagellar segments with dense yellowish short setae. Thorax. Scutum densely covered with setae, with marginal and prescutellar setae stronger. Antepronotum with 6–7 strong and a number of weaker setae. Proepisternum with one very strong seta at anterior margin about 10 weaker setae. Anepisternum, katepisternum and anepimeron non-setose. Laterotergite with long fine setae on posterior half. Mediotergite non-setose. Scutellum with a row of marginal setae including two pairs remarkably stronger. Wing. Length 5.3–5.7 mm, length to width 2.3–2.8. All veins setose, except Sc, sc-r, Rs and extreme base of M1. Sc ending on C at level of furcation of posterior fork. R4+5 3.3 times as long as R1. r-m 1.47 times as long as m-stem. M1 and M2 apically convergent, apical third of both veins faint. M4 interrupted at base. Rs located distally of the anterior fork. Legs. Ratio of femur to tibia for fore, mid and hind legs: 1.16; 0.95; 0.86. Ratio of tibia to basitarsus for fore, mid and hind legs: 1.00; 1.66; 2.00. Fore tibia with a spur 2.95 times of tibial maximum width. Mid tibia with anterior spur 3.33 times and posterior spur 3.96 times of tibial maximum width. Hind tibia with anterior spur 3.33 times and posterior spur 4.58 times of tibial maximum width. Terminalia (Figs 5A–D, 6A–C). Gonocoxite with setae on apical fifth only. Ventromedial process of gonocoxite elongated ovoid with a row of long setae apically and an aggregation of shorter setae on apical fourth. Ventroposterior margin of gonocoxite drawn into a wide membraneous non-setose medial lobe and a digitate more protruding lobe with one prominent and 2–3 weaker apical setae. Tergite 9 membraneous, somewhat tapering, apically evenly rounded with apicocentral patch of short setae. Gonostylus bipartite: lateral prong shorter, convolute and apically hooked; medial prong longer, tapering with preapical small tooth at ventral margin. Aedeagus with sclerotized, cup-shaped apical portion, ejaculatory apodeme bilobed. Paramere about 1.6 times as long as aedeagus, bowed in lateral view, apically tapering, with ventral flange drawn out into a triangular membranous process in the middle; anteriorly, parameres fused into a complex membranous structure with anterior concavity and well protruding lateral corners. Hypoproct with protruding apicolateral corners and medial part that bears a group of stout setae.

Figure 5. 

Leia katae sp. nov., male terminalia A dorsal view B lateral view C ventral view D gonocoxite and gonostylus, ventral view. Abbreviations: aed = aedeagus, cer = cercus, ej ap = ejaculatory apodeme, gc = gonocoxite, gc vmp = ventromedial process of gonocoxite, gc vpp = ventroposterior process of gonocoxite, gst lp = lateral prong of gonostylus, gst mp = medial prong of gonostylus, hyp = hypoproct, par = paramere, tg 9 = tergite IX. Scale bars: 0.2 mm.

Female. Unknown.

Etymology

The species is named after my daughter Katariina (born 3 November 2000), an enthusiastic student of biology at the University of Tartu (Estonia). She participated in a trip to Georgia in 2017 that yielded several specimens of this study and she always insists we call her Kata.

Remarks

There are 166 Leia Meigen species known worldwide including 33 in the Palaearctic region (Polevoi and Salmela 2016). Leia katae sp. nov. differs from all known Palaearctic and Nearctic species by its peculiar structure of the gonostylus that is bipartite: medial prong long and slender with a preapical tooth, and lateral prong apically hooked, about 2/3 of the medial prong.

Figure 6. 

Leia katae sp. nov., male terminalia A aedeagal complex, ventral view B aedeagal complex, lateral view C hypoproct ventral view. Abbreviations: aed = aedeagus, ej ap = ejaculatory apodeme, par = paramere. Scale bars: 0.2 mm.

Anatella metae sp. nov.

http://zoobank.org/7CD91319-0672-4DC8-9CF6-7140ADF1F13E
Figs 3B, 7A–G

Type material

Holotype. Male, Georgia, Mtskhetha-Mthianethi, Saguramo north of Tbilisi, 41°53.07'N, 44°46.78'E, 920 m a.s.l., 15.v.2013, sweeping, leg. O. Kurina [see Table 1: MM-12] (mounted from alcohol, IZBE). Paratype. Male, same as holotype (mounted from alcohol, IZBE).

Diagnosis

Anatella metae sp. nov. is characterized by the presence of a strong posteroventral fringe on mid femora with a row of strong setae, absence of anterior spur on mid tibia, absence of setae on hind coxa basally. The new species is closest to A. atlanticiliata Chandler and Ribeiro but differs in characters of the male terminalia: ventral branch of the gonostylus about twice as long as the dorsal branch, dorsal branch of the gonostylus with long and slender medial prong, medial branch of the gonostylus slender and apically hooked.

Description

Male. Body length 2.7–2.9 mm (n = 2). Coloration. Head with vertex, frons, face and clypeus brown, mouthparts including palpus pale yellow. Scape, pedicel and base of first flagellomere yellow, rest of flagellum light brown. Thorax with scutum and lateral parts light brown. Thoracic setae yellowish to brown, with thicker setae darker than finer ones. Wing hyaline, unmarked with yellowish tinge. Halter with stem and knob pale yellow. Legs yellow, tarsi yellow but seem darker because of dense brown setae. All setae on legs brown, tibial spurs yellowish. Abdomen mainly brown with first two segments somewhat lighter. Abdominal vestiture brown. Terminalia light brown. Head. Ocelli two, very close to eye margins, with dark brown patches at anterior margin. Frontal furrow complete. Clypeus rectangular. Fourth flagellar segment about 2.5 times as long as wide, apical flagellar segment 2.5 times as long as wide basally. Flagellar segments with dense whitish setae about one third of segments’ width. Thorax. Scutum covered with setae, with marginal and prescutellar setae stronger. Antepronotum with 2 strong and 10–15 weaker setae. Proepisternum with two strong and 2–3 weaker setae. Anepisternum, katepisternum and anepimeron non-setose. Laterotergite with about 10 setae on upper half. Mediotergite non-setose. Scutellum with about 10 setae on upper surface. Wing. Length 2.39–2.70 mm, length to width 2.75–2.90. C, R, R1, R4+5 setose, all other veins non-setose. C produced halfway between R4+5 and M1. r-m about as long as m-stem. Posterior fork at the level of anterior fork or slightly before. CuA slightly sinuous. Legs. Ratio of femur to tibia for fore, mid and hind legs: 1.08–1.17; 0.97–1.00; 0.65–0.90. Ratio of tibia to basitarsus for fore, mid and hind legs: 0.96–1.00; 1.21–1.28; 1.41–1.77. Fore tibia with a spur 2.00 times of tibial maximum width. Mid tibia with anterior spur absent and posterior spur 2.27–2.40 times of tibial maximum width. Hind tibia with anterior spur 2.71–3.33 times and posterior spur 3.93–4.66 times of tibial maximum width. Strong posteroventral fringe of mid femora with row of strong setae. Hind coxa without basal setae. Terminalia (Fig. 7A–G). Gonocoxite ventrally with (1) V-shaped wide incision anteriorly, (2) posteromedial non-setose tapering projection with deep slit, and (3) posterolateral large apically setose lobes. Gonostylus divided into four branches (Fig. 7F–G). The ventral branch of the gonostylus elongated digitate, apical half setose and with one strong seta apically deviating from other setosity. Dorsal branch of the gonostylus about half length of the ventral branch, divided into two prongs: medial finger like bare prong and lateral large apically and basally setose prong. Medial branch of the gonostylus slightly shorter than dorsal branch, slender, apically hooked. Internal branch of the gonostylus membranous, convolute with anterior lamellae.

Figure 7. 

Anatella metae sp. nov. male terminalia A dorsal view B ventral view C dorsal view, cerci and tergite IX removed D lateral view E cerci and tergite IX, dorsal view F, G gonostylus, internal views from different angles. Abbreviations: aed complex – aedeagal complex, cer = cercus, gc = gonocoxite, gc ap = gonocoxal apodeme, gst dbr = dorsal branch of gonostylus, gst ibr = internal branch of gonostylus, gst mbr = medial branch of gonostylus, gst vbr = ventral branch of gonostylus, tg 9 = tergite IX. Scale bars: 0.1 mm.

Etymology

The species is named after my daughter Liisa-Meta (born 9 October 2004), a keen naturalist who also participated in a trip to Georgia in 2017.

Remarks

There are about 50 Anatella Winnertz species known in the Holarctic region (cf. Fungus Gnats Online Authors 2021), the vast majority of which are adequately described and figured. In having posteroventral fringe of mid femora with strong setae and absence of anterior spur of mid tibia, A. metae sp. nov. resembles A. atlanticiliata Chandler & Ribeiro, 1995 known only from Madeira Island. Also, the male terminalia of both species share the general outline (cf. Chandler and Ribeiro 1995: fig. 27). However, A. metae differs in the structure of the gonostylus as follows: (1) ventral branch of the gonostylus twice as long as dorsal branch (only somewhat longer in A. atlanticiliata), (2) dorsal branch of the gonostylus with medial prong long, anchored to lateral prong basally (medial prong short, anchored to lateral prong subapically in A. atlanticiliata), and (3) medial branch of gonostylus slender, apically hooked (medial branch stout, slightly curved in A. atlanticiliata).

List of fungus gnat species of Georgia

Family Bolitophilidae

Bolitophila (Bolitophila) austriaca (Mayer, 1950)

Material

2♀♀, SJ-7; 4♀♀, SJ-8. Total: 6♀♀.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Bolitophila (Bolitophila) basicornis (Mayer, 1951)

Fig. 9A

Material

1♀, MM-1. Total: 1♀.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Bolitophila (Bolitophila) cinerea Meigen, 1818

Material

1♂, MM-12. Total: 1♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Bolitophila (Cliopisa) fumida Edwards, 1941

Material

1♂, SJ-9. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Family Diadocidiidae

Diadocidia (Adidocidia) valida Mik, 1874

Material

1♀, SJ-1; 1♂, SK-1. Total: 1♂ 1♀.

Distribution in Georgia

Shida Kartli, Samtskhe-Javakheti.

General distribution

Western Palaearctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 1994).

Diadocidia (Diadocidia) ferruginosa (Meigen, 1830)

Material

1♂, SZS-3 (ZFMK); 2♂♂ 1♀, I-6; 2♂♂, I-9; 1♂, I-11; 1♂, A-1; 1♂, SJ-4; 1♂, SJ-7; 1♂, SJ-8; 4♂♂, SK-1. Total: 14♂♂ 1♀.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Shida Kartli, Imereti, Samtskhe-Javakheti.

General distribution

Holarctic.

Diadocidia (Diadocidia) spinosula Tollet, 1948

Fig. 8G

Material

1♂, SZS-3 (IZBE); 2♂♂, SK-1; 2♂♂, SJ-1 (ZFMK); 1♂, SJ-2 (IUTG); 1♂ 1♀, SJ-4; 2♂♂, SJ-7; 4♂♂ 3♀♀, SJ-9. Total: 13♂♂ 4♀♀.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Shida Kartli, Samtskhe-Javakheti.

General distribution

Palaearctic.

Figure 8. 

Habitus of Georgian fungus gnats of the families Ditomyiidae (A), Keroplatidae (B–F) and Diadocidiidae (G) A Symmerus annulatus (Meigen, 1830) B Keroplatus testaceus Dalman, 1818 C Macrorrhyncha flava Winnertz, 1846 D Pyratula zonata (Zetterstedt, 1855) E Cerotelion racovitzai Matile & Burghele-Balacesco, 1969 F Macrocera vittata Meigen, 1830 G Diadocidia (Diadocidia) spinosula Tollet, 1948.

Family Ditomyiidae

Ditomyia fasciata (Meigen, 1818)

Material

2♂♂, I-9. Total: 2♂♂.

Distribution in Georgia

Imereti.

General distribution

Palaearctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 1994).

Symmerus annulatus (Meigen, 1830)

Fig. 8A

Material

6♂♂, SZS-3 (2♂♂ ZFMK, 2♂♂ IUTG, 2♂♂ IZBE); 1♂, A-1; 1♂ 1♀, A-3; 1♀, I-3 (18.v–1.vi.2013); 1♂, I-6; 1♂ 1♀, I-9; 1♂, I-10; 2♂♂, I-11; 1♂, I-12; 4♂♂, SJ-1 (2♂♂ ZFMK, 1♂ IUTG, 1♂ IZBE); 2♂♂, MM-12. Total: 20♂♂ 3♀♀.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Adjara, Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 1994).

Family Keroplatidae

Subfamily Macrocerinae

Macrocera centralis Meigen, 1818

Material

2♂♂, KK-1; 1♀, MM-14. Total: 2♂♂ 1♀.

Distribution in Georgia

Kvemo Kartli, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Macrocera crassicornis Winnertz, 1863

Material

1♂, A-1. Total: 1♂.

Distribution in Georgia

Adjara.

General distribution

Palaearctic.

Remarks

In Transcaucasia recorded from Armenia (Zaitzev 1994).

Macrocera fasciata Meigen, 1804

Material

1♂, SZS-3 (ZFMK). Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Palaearctic.

Macrocera fastuosa Loew, 1869

Material

2♂♂, A-1; 3♂♂, A-7. Total: 5♂♂.

Distribution in Georgia

Adjara.

General distribution

Europe.

Macrocera lutea Meigen, 1804

Material

1 ♂, A-7; 1♂, KK-1. Total: 2♂♂.

Distribution in Georgia

Adjara, Kvemo Kartli.

General distribution

Palaearctic.

Remarks

In Transcaucasia recorded from Armenia (Joost and Plassmann 1985).

Macrocera phalerata Meigen, 1818

Material

1♀, A-1; 1♀, A-6; 1♀, A-7. Total: 3♀♀.

Distribution in Georgia

Adjara.

General distribution

Palaearctic.

Macrocera stigma Curtis, 1837

Material

1♂ 1♀, A-1; 4♂♂, A-7; 11♂♂ 3 ♀♀, I-6; 2♂♂, I-11; Total: 18♂♂ 8♀♀.

Distribution in Georgia

Adjara, Imereti.

General distribution

Palaearctic.

Macrocera stigmoides Edwards, 1925

Material

44♂♂ 14♀♀, KK-1. Total: 44♂♂ 14♀♀.

Distribution in Georgia

Kvemo Kartli.

General distribution

Palaearctic.

Macrocera vittata Meigen, 1830

Fig. 8F

Material

3♂♂, SJ-8; 2♂♂, MM-13; 1♂, MM-14. Total: 6♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Subfamily Keroplatinae

Tribe Keroplatini

Cerotelion racovitzai Matile & Burghele-Balacesco, 1969

Fig. 8E

Material

6♂♂, A-3; 1♂, A-5; 3♂♂, A-7; 14♂♂ 1♀, I-6; 1♀, I-8; 3♂♂, I-9; 1♂, I-10; 2♂♂, I-11; 2♂♂, I-14; 1♂, I-15; 1♀, MM-7; 3♂♂, MM-8. Total: 36♂♂ 3♀♀.

Distribution in Georgia

Adjara, Imereti, Mtskhetha-Mthianethi.

General distribution

Western Palaearctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 1994).

Cerotelion striatum (Gmelin, 1790)

Material

1♂, I-4. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Western Palaearctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 1994).

Keroplatus testaceus Dalman, 1818

Fig. 8B

Material

2♂♂, I-6. Total: 2♂♂.

Distribution in Georgia

Imereti.

General distribution

Palaearctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 1994).

Tribe Orfeliini

Isoneuromyia semirufa (Meigen, 1818)

Georgian source

Zaitzev 1994: 82 (from Adjara).

Distribution in Georgia

Adjara.

General distribution

Holarctic.

Remarks

Zaitzev (1994) studied a single male specimen from Batumi collected in 1908. The black colour of the body as noted by Zaitzev (1994) for the studied material is characteristic to I. semirufa. The other European species have the thorax yellow to orange with or without longitudinal stripes; also, see the next species and discussion by Mantič and Ševčík (2017).

Isoneuromyia czernyi (Strobl, 1909)

Material

1♂, A-1; 1♂, A-5; 1♀, A-7; 1♂ 1♀, I-6. Total: 3♂♂ 2♀♀.

Distribution in Georgia

Adjara, Imereti.

General distribution

Europe.

Remarks

All studied Georgian specimens correspond to the diagnosis including figures provided recently by Mantič and Ševčík (2017), i.e. (1) scutum with longitudinal dark stripes which are, however, almost fused in female specimens (thorax all dark brown to blackish in I. semirufa), (2) wing with a distinct subapical band (anteriorly infuscated in I. semirufa) and (3) male terminalia with medial tooth of the gonostylys larger than the lateral one (both in subequal size in I. semirufa). I. czernyi is a rare European species known from the Mediterranean region and Slovakia (Mantič and Ševčík 2017).

Macrorrhyncha flava Winnertz, 1846

Fig. 8C

Material

2♂♂, K-4; 7♂♂ 3♀♀, K-5. Total: 9♂♂ 3♀♀.

Distribution in Georgia

Kakheti.

General distribution

Europe.

Monocentrota lundstromi Edwards, 1925

Material

1♂, SZS-3 (IZBE). Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Europe.

Neoplatyura modesta (Winnertz, 1863)

Material

1♂, K-2; 1♀, K-3. Total: 1♂ 1♀.

Distribution in Georgia

Kakheti.

General distribution

Europe.

Neoplatyura nigricauda (Strobl, 1893)

Material

1♂, SZS-1. Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Europe.

Orfelia discoloria (Meigen, 1818)

Material

7♂♂, SZS-3(2♂♂ ZFMK, 3♂♂ IUTG, 2♂♂ IZBE); 1♂, A-3; 1♂, SJ-1 (ZFMK); 1♂, MM-7. Total: 10♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Adjara, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Holarctic.

Orfelia georgica Kurina & Jürgenstein, 2013

Fig. 9B

Georgian source

Kurina and Jürgenstein 2013: 23 (fig. 2a–d)

Type material

1♂, I-10 (holotype); 1♂, I-9 (paratype); 10♂♂, I-6 (paratypes). Additional material. 12♂♂ 4♀♀, A-1; 34♂♂, A-3; 7♂♂, A-5; 76♂♂, A-7; 1♂, A-8; 5♂♂, I-6; 12♂♂, I-11; 1♂, I-12; 5♂♂, SJ-1; 1♂, SJ-2 (ZFMK); 4♂♂, K-4; 1♂, K-6. Total: 171♂♂ 4♀♀.

Distribution in Georgia

Adjara, Imereti, Samtskhe-Javakheti, Kakheti.

General distribution

Georgia.

Figure 9. 

Habitus of Georgian fungus gnats of the families Bolitophilidae (A), Keroplatidae (B) and Mycetophilidae (C–I) A Bolitophila (Bolitophila) basicornis (Mayer, 1951) B Orfelia georgica Kurina & Jürgenstein, 2013 C Acnemia nitidicollis (Meigen, 1818) D Monoclona rufilatera (Walker, 1837) E Azana (Azana) anomala (Staeger, 1840) F Phthinia hyrcanica Zaitzev, 1984 G Neoempheria striata (Meigen, 1818) H Neoempheria brevilineata Okada, 1939 I Mycomya (Mycomya) marginata (Meigen, 1818).

Orfelia trifida Kurina & Jürgenstein, 2013

Georgian source

Kurina and Jürgenstein 2013: 24 (fig. 3a–d).

Type material

1♂, I-5 (holotype). Additional material. 35♂♂, SZS-3 (12♂♂ ZFMK, 12♂♂ IUTG, 11♂♂ IZBE). Total: 36♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti.

General distribution

Georgia.

Pyratula perpusilla (Edwards, 1913)

Material

16♂♂, SJ-4. Total: 16♂♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Europe.

Remarks

The P. perpusilla species-group includes at least seven closely related species in Europe, separable only by small details of male terminalia (Chandler and Blasco-Zumeta 2001). The studied Georgian specimens have the ventroapical margin of the gonocoxite with setose lobe (= without asetose protuberance) that is shared by three species, viz. P. perpusilla, P. alpicola Chandler, 2001 and P. oracula Chandler, 1994. The aedeagal complex is considerably short (elongate in P. oracula) and the aedeagal sheath is interrupted medially on the ventral side (with complete bridge in P. alpicola). However, the Georgian specimens are slightly different from P. perpusilla as figured by Chandler and Blasco-Zumeta (2001: Figs 9–12) in having the distal dorsal corner of the aedeagal seath with a blunt protuberance that is otherwise typical to P. alpicola. The material was compared to that of P. alpicola and P. oracula from North Italy (cf. Kurina 2008b) and, pending a further molecular study of this species-group, is considered to be conspecific with P. perpusilla.

Pyratula zonata (Zetterstedt, 1855)

Fig. 8D

Material

2♂♂, A-5; 1♂, I-6; 2♂♂, I-11; 2♂♂, SJ-1 (1♂ IUTG, 1♂ IZBE); 1♂ 1♀, SJ-2 (ZFMK); 1♂, SJ-3; 4♂♂, MM-11. Total: 13♂♂ 1♀.

Distribution in Georgia

Adjara, Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Europe.

Urytalpa dorsalis (Staeger, 1840)

Material

1♂, SZS-3 (IZBE); 3♂♂, SJ-1 (1♂ ZFMK, 1♂ IUTG, 1♂ IZBE); 1♂ 2♀♀, SJ-2 (1♂ 1♀ ZFMK, 1♀ IUTG). Total: 5♂♂ 2♀♀.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Samtskhe-Javakheti.

General distribution

Europe.

Subfamily Platyurinae

Platyura marginata Meigen 1804

Material

1♂, K-6. Total: 1♂.

Distribution in Georgia

Kakheti.

General distribution

Palaearctic.

Family Mycetophilidae

Subfamily Mycomyinae

Mycomya (Cymomya) circumdata (Staeger, 1840)

Material

2♂♂, SZS-3 (1♂ ZFMK, 1♂ IZBE). Total: 2♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Palaearctic.

Mycomya (Neomycomya) fimbriata (Meigen, 1818)

Material

1♂, A-7. Total: 1♂.

Distribution in Georgia

Adjara.

General distribution

Holarctic, extending to the Oriental region.

Mycomya (Mycomya) bialorussica Landrock, 1925

Material

1♂, SZS-3 (ZFMK). Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Europe.

Mycomya (Mycomya) cinerascens (Macquart, 1826)

Material

1♂, SJ-8. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Holarctic, extending to the Oriental region.

Mycomya (Mycomya) flavicollis (Zetterstedt, 1852)

Material

6♂♂, A-5; 1♂, A-7; 4♂♂, I-6; 2♂♂, SJ-3; 29♂♂, SJ-4; 1♂, SJ-9; 1♂, MM-7; 2♂♂, MM-14; 1♂, K-6. Total: 47♂♂.

Distribution in Georgia

Adjara, Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi, Kakheti.

General distribution

Western Palaearctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 1994).

Mycomya (Mycomya) griseovittata (Zetterstedt, 1852)

Material

1♂, SZS-3 (ZFMK). Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Holarctic.

Mycomya (Mycomya) marginata (Meigen, 1818)

Fig. 9I

Material

3♂♂, I-6; 1♂, I-17; 1♂, SJ-3; 2♂♂, MM-8; 1♂, MM-12. Total: 8♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Mycomya (Mycomya) occultans (Winnertz, 1863)

Material

1♂, SJ-4. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic, extending to the Oriental region.

Mycomya (Mycomya) tenuis (Walker, 1856)

Material

1♂, A-7; 1♂, SJ-4. Total: 2♂♂.

Distribution in Georgia

Adjara, Samtskhe-Javakheti.

General distribution

Palaearctic.

Mycomya (Mycomya) tridens (Lundström, 1911)

Material

1♂, SZS-4 (ZFMK). Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Europe.

Mycomya (Mycomya) tumida (Winnertz, 1863)

Material

1♂, SZS-4 (ZFMK); 1♂, I-11. Total: 2♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti.

General distribution

Palaearctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 1994).

Mycomya (Mycomya) winnertzi (Dziedzicki, 1885)

Material

1♂, SK-1; 1♂, SJ-8. Total: 2♂♂.

Distribution in Georgia

Shida Kartli, Samtskhe-Javakheti.

General distribution

Palaearctic, extending to the Oriental region.

Mycomya (Mycomyopsis) affinis (Staeger, 1840)

Material

3♂♂, K-5; 1♂, K-6. Total: 4♂♂.

Distribution in Georgia

Kakheti.

General distribution

Palaearctic.

Mycomya (Mycomyopsis) trilineata (Zetterstedt, 1838)

Material

5♂♂, K-6. Total: 5♂♂.

Distribution in Georgia

Kakheti.

General distribution

Palaearctic.

Neoempheria brevilineata Okada, 1939

Figs 9H, 10A–F

Material

1♂, A-7; 2♂♂, I-6. Total: 3♂♂.

Distribution in Georgia

Adjara, Imereti.

General distribution

Palaearctic.

Remarks

The species description from Hokkaido (Japan) by Okada was supplemented by a black and white figure of the general habitus including wing venation and pattern (Okada 1939: plate XVI, fig. 3). The Georgian material was compared to that from Japan (1 ♂, JAPAN, Honshu, Ishikawa Perfecture, Kanazawa City, Kakuma Campus, window trap, 14.vii-21.vii.2006, Indah, T. leg.; Kjærandsen J. det., TSZD-JKJ-111335) and the small differences in male terminalia are considered to be within intraspecific variation. Figures of the male terminalia (Fig. 10A–F) are provided for the first time for the species.

Figure 10. 

Neoempheria brevilineata Okada, 1939, male terminalia A, B, dorsal view C, D ventral view E lateral view F posterior view. Scale basr: 0.2 mm.

Neoempheria striata (Meigen, 1818)

Fig. 9G

Material

1♀, I-6; 1♂ 1♀, I-14. Total: 1♂ 2♀♀.

Distribution in Georgia

Imereti.

General distribution

Palaearctic.

Subfamily Sciophilinae

Acnemia amoena Winnertz, 1863

Material

1♂, I-6. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Palaearctic.

Acnemia hyrcanica Zaitzev, 1984

Material

1♂, SZS-3 (ZFMK); 1♂, I-6. Total: 2♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti.

General distribution

Caucasia.

Remarks

Recorded earlier from North Caucasus and Azerbaijan (Zaitzev 1994).

Acnemia nitidicollis (Meigen, 1818)

Fig. 9C

Material

4♂♂, SZS-3 (2♂♂ ZFMK, 1♂ IUTG, 1♂ IZBE); 2♂♂, A-3; 2♂♂, A-7; 3♂♂ 1♀, K-4; 1♂, K-5; 1♂, K-6. Total: 13♂♂ 1♀.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Adjara, Kakheti.

General distribution

Palaearctic.

Allocotocera pulchella (Curtis, 1837)

Fig. 11H

Material

1♂, SJ-2 (ZFMK). Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Holarctic.

Figure 11. 

Habitus of Georgian fungus gnats of the family Mycetophilidae A Boletina trivittata (Meigen, 1818) B Synapha fasciata Meigen, 1818 C Grzegorzekia collaris (Meigen, 1818) D Clastobasis loici Chandler, 2001 E Manota unifurcata Lundström, 1913 F Megalopelma nigroclavatum (Strobl, 1910) G Leia winthemii Lehmann, 1822 H Allocotocera pulchella (Curtis, 1837) I Coelosia flava (Staeger, 1840) J Novakia scatopsiformis Strobl, 1893 K Ectrepesthoneura hirta (Winnertz, 1846).

Anaclileia adjarica Kurina, 2018

Georgian source

Kurina 2018: 156 (figs 2–5).

Type material

1♂, A-1 (holotype); 3♂♂, A-1 (paratypes); 3♂♂ 3♀♀, A-3 (paratypes); 1♂, A-7 (paratype). Total: 8♂♂ 3♀♀.

Distribution in Georgia

Adjara.

General distribution

Georgia.

Remarks

The species was recently described from material collected from Mtirala and Kintrishi National Parks in Adjara (Kurina 2018)

Azana (Azana) anomala (Staeger, 1840)

Fig. 9E

Material

1♀, SJ-3. Total: 1♀.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Europe.

Azana (Jugazana) nigricoxa Strobl, 1898

Material

1♂, I-11. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Europe.

Megalopelma nigroclavatum (Strobl, 1910)

Fig. 11F

Material

2♂♂, I-6. Total: 2♂♂.

Distribution in Georgia

Imereti.

General distribution

Holarctic.

Monoclona rufilatera (Walker, 1837)

Fig. 9D

Material

3♂♂, I-6; 1♂, SK-1. Total: 4♂♂.

Distribution in Georgia

Imereti, Shida Kartli.

General distribution

Holarctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 1994).

Neuratelia caucasica Zaitzev, 1994

Georgian source

Kurina et al. 2015: 116 (figs 11, 12, 16).

Material

1♀, SZS-3 (IZBE); 2♀♀, SZS-4 (1♀ ZFMK, 1♀ IUTG); 3♂♂ 4♀♀, A-3; 1♂ 2♀♀, A-7; 2♂♂ 10♀♀, I-6; 1♂ 1♀, I-9; 2♂♂ 2♀♀, SK-1; 1♂, SJ-1 (IZBE); 4♂♂ 1♀, SJ-2 (2♂♂ ZFMK, 1♂ 1♀ IUTG, 1♂ IZBE); 2♂♂ 1♀, SJ-3; 1♂, KK-1. Total: 17♂♂ 24♀♀.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Adjara, Imereti, Shida Kartli, Samtskhe-Javakheti, Kvemo Kartli.

General distribution

Caucasia: Russia (Krasnodarskiy region), Georgia.

Phthinia hyrcanica Zaitzev, 1984

Fig. 9F

Material

1♂, MM-8; 1♂, K-4. Total: 2♂♂.

Distribution in Georgia

Mtskhetha-Mthianethi, Kakheti.

General distribution

Caucasia.

Remarks

Known only from type locality in Azerbaijan (Zaitzev 1994).

Polylepta zonata Zetterstedt, 1852

Material

1♂, A-5. Total: 1♂.

Distribution in Georgia

Adjara.

General distribution

Europe, with scattered distribution (Kurina 2003, Chandler 2013).

Sciophila fenestella Curtis, 1837

Material

1♂, I-6. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Holarctic.

Sciophila georgei sp. nov.

Material

See in species description above.

Distribution in Georgia

Kakheti.

General distribution

Georgia.

Sciophila nitens (Winnertz, 1863)

Material

1♂, SZS-3 (ZFMK); 1♂, SJ-1 (IZBE); 1♂, SJ-2 (IUTG). Total: 3♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Samtskhe-Javakheti.

General distribution

Holarctic.

Remarks

In Europe recorded from mountain areas (Kurina 2004, 2008b).

Sciophila thoracica Staeger, 1840

Material

1♂, SZS-3 (IZBE); 1♂, SJ-2 (ZFMK). Total: 2♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Samtskhe-Javakheti.

General distribution

Holarctic.

Syntemna morosa Winnertz, 1863

Material

1♂, I-6; 1♂, I-9. Total: 2♂♂.

Distribution in Georgia

Imereti.

General distribution

Europe.

Subfamily Gnoristinae

Apolephthisa subincana (Curtis, 1837)

Material

1♂, I-6. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Western Palaearctic.

Boletina borealis Zetterstedt, 1852

Material

1♂, SZS-2; 1♂, SZS-4 (ZFMK). Total: 2♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Palaearctic.

Boletina digitata Lundström, 1914

Material

4♂♂, SZS-4 (2♂♂ ZFMK, 1♂ IUTG, 1♂ IZBE). Total: 4♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Western Palaearctic.

Boletina dubia (Meigen, 1804)

Material

5♂♂, A-1; 3♂♂, A-3. Total: 8♂♂.

Distribution in Georgia

Adjara.

General distribution

Europe.

Remarks

The Georgian specimens have the ventral lobe of the gonostylus with a blunt small spine apically that is absent in studied specimens from Estonia and Sweden as well as in published figures (e.g. Landrock 1927, Hutson et al. 1980, Zaitzev 1994). Otherwise, the male terminalia including aedeagal complex do not have any substantial differences. Therefore, the Georgian material is considered to be conspecific pending further, more thorough study including that based on DNA sequencing.

Boletina gripha Dziedzicki, 1885

Material

2♂♂, SZS-4 (1♂ ZFMK, 1♂ IZBE). Total: 2♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Palaearctic.

Boletina moravica Landrock, 1912

Material

1♂, SZS-4 (ZFMK). Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Europe.

Boletina nitida Grzegorzek, 1885

Material

10♂♂, SZS-3 (4♂♂ ZFMK, 4 ♂♂ IUTG, 2 ♂♂ IZBE); 1♂, SJ-1 (IZBE); 1♂, K-6. Total: 12♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Samtskhe-Javakheti, Kakheti.

General distribution

Palaearctic.

Boletina sciarina Staeger, 1840

Material

3♂♂, SZS-4 (1♂ ZFMK, 1♂ IUTG, 1♂ IZBE). Total: 3♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Holarctic.

Boletina trivittata (Meigen, 1818)

Fig. 11A

Material

4♂♂, SJ-8; 5♂♂, SJ-9. Total: 9♂♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Coelosia flava (Staeger, 1840)

Fig. 11I

Georgian source

Thormann et al. 2019: 279 (from Mtskhetha-Mthianethi).

Material

1♂, SZS-3 (ZFMK); 8♂♂ 1♀, KK-1. Total: 9♂♂ 1♀.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Mtskhetha-Mthianethi, Kvemo Kartli.

General distribution

Europe.

Docosia gilvipes (Haliday in Walker, 1856)

Georgian source

Ševčík et al. 2020: 21

Material

1♂, SZS-3 (ZFMK); 1♀, I-6. Total: 1♂ 1♀.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti.

General distribution

Palaearctic.

Docosia flavicoxa Strobl, 1900

Georgian source

Ševčík et al. 2020: 21

Material

3♂♂, K-5. Total: 3♂♂.

Distribution in Georgia

Kakheti.

General distribution

Europe.

Docosia moravica Landrock, 1916

Georgian source

Ševčík et al. 2020: 23.

Material

1♂, SJ-2 (ZFMK). Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Docosia pannonica Laštovka & Ševčík, 2006

Georgian source

Ševčík et al. 2020: 23

Material

1♂, MM-3. Total: 1♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Europe.

Remarks

Known from Central Europe (Laštovka and Ševčík 2006).

Docosia svanetica Kurina in Ševčík et al. 2020

Georgian source

Ševčík et al. 2020: 17.

Material

1♂, SZS-4 (holotype, ZFMK); 5♂♂, SZS-4; 2♂♂, SZS-3; 1♂, SJ-1. Total: 9♂♂ (see Ševčík et al. 2020 for depository of paratypes).

Distribution in Georgia

Samegrelo-Zemo Svanethi, Samtskhe-Javakheti.

General distribution

Georgia.

Ectrepesthoneura hirta (Winnertz, 1846)

Fig. 11K

Material

1♂, SJ-2 (ZFMK); 1♂, K-6. Total: 2♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Kakheti.

General distribution

Europe.

Grzegorzekia collaris (Meigen, 1818)

Fig. 11C

Georgian source

Thormann et al. 2019: 279 (from Mtskhetha-Mthianethi).

Material

2♂♂, SZS-3 (1♂ ZFMK, 1♂ IZBE); 1♂, I-10; 1♂, SK-1. Total: 4♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Shida Kartli, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Lusitanoneura chandleri (Caspers, 1991)

Material

1♂, SZS-3 (ZFMK); 2♂♂, I-6. Total: 3♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti.

General distribution

Europe.

Remarks

Known only from Grete and Cyprus (Caspers 1991, Chandler et al. 2006, Ribeiro and Chandler 2007).

Palaeodocosia vittata (Coquillett, 1901)

Material

1♂, A-1; 1♂, I-6. Total: 2♂♂.

Distribution in Georgia

Adjara, Imereti.

General distribution

Holarctic.

Synapha fasciata Meigen, 1818

Fig. 11B

Material

104♂♂, A-1; 54♂♂ 5♀♀, A-2; 70♂♂, A-3; 21♂♂ 2♀♀, A-4; 6♂♂ 4♀♀, A-5; 3♂♂, A-6; 173♂♂ 64♀♀, A-7; 14♂♂, I-1; 7♂♂, I-3 (18.v–1.vi.2013); 1♂, I-4; 138♂♂ 7♀♀, I-6; 25♂♂, I-7; 2♂♂ 3♀♀, I-9; 1♀, I-10; 8♂♂, I-11; 15♂♂, I-13; 2♂♂, I-16; 37♂♂, I-17; 1♂, MM-9. Total: 683♂♂ 86♀♀.

Distribution in Georgia

Adjara, Imereti, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Subfamily Leiinae

Clastobasis alternans (Winnertz, 1863)

Material

6♂♂ 2♀♀, SZS-3 (2♂♂ 1♀ ZFMK, 2♂♂ 1♀ IUTG, 2♂♂ IZBE); 1♂, K-4; 1♂, K-5. Total: 8♂♂ 2♀♀.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Kakheti.

General distribution

Palaearctic.

Clastobasis loici Chandler, 2001

Fig. 11D

Material

3♂♂, K-4; 22♂♂, K-5. Total: 25♂♂.

Distribution in Georgia

Kakheti.

General distribution

Palaearctic.

Remarks

This very rare species was until recently recorded only from Channel Islands and Central Europe but Kurina (2020b) found it also from Japan. The record from Georgia suggests a wider distribution in the Palearctic region.

Greenomyia mongolica Laštovka & Matile, 1974

Material

1♂, SJ-5. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Remarks

A widely distributed Palaearctic species that has expanded its range in Europe during recent decades and is locally common also in anthropogenic environments (Kurina et al. 2011, pers. observations).

Leia bimaculata (Meigen, 1804)

Material

1♂, SZS-3 (IZBE); 1♂, SK-1; 1♂, SJ-2 (ZFMK); 1♂, SJ-9; 1♂, MM-7. Total: 5♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Shida Kartli, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Remarks

Aedeagal complex of Georgian specimens is similar to that figured by Polevoi and Salmela (2016: fig. 7G) and gonostylus has a clear dorsal projection (Polevoi and Salmela 2016: fig. 7I, J).

Leia cylindrica (Winnertz, 1863)

Material

1♂ 1♀, I-6; 1♂, K-6. Total: 2♂♂ 1♀.

Distribution in Georgia

Imereti, Kakheti.

General distribution

Western Palaearctic.

Leia katae sp. nov.

Material

See in species description above.

Distribution in Georgia

Samegrelo-Zemo-Svanethi, Shida-Kartli.

General distribution

Georgia.

Leia piffardi Edwards, 1925

Material

1♂, I-6. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Europe, with scattered distribution.

Leia winthemii Lehmann, 1822

Fig. 11G

Material

3♂♂, I-10; 1♂, SJ-8. Total: 4♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti.

General distribution

Holarctic, extending to the Oriental region.

Novakia scatopsiformis Strobl, 1893

Fig. 11J

Material

1♂ 1♀, SJ-2 (1♀ ZFMK, 1♂ IZBE); 1♂, K-4; 1♀, K-5. Total: 2♂♂ 2♀♀.

Distribution in Georgia

Samtskhe-Javakheti, Kakheti.

General distribution

Western Palaearctic.

Remarks

According to the recent molecular study by Kaspřák et al. (2019: Fig. 1), the genus Novakia Strobl, 1893 apparently belongs to the subfamily Gnoristinae. However, as the authors did not have further discussion about this relationship, I follow the current classification in Fungus Gnats Online (http://www.sciaroidea.info/).

Subfamily Manotinae

Manota unifurcata Lundström, 1913

Fig. 11E

Material

1♂, I-6. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Europe.

Remarks

A rare species, recorded from Central and Northern Europe with the south-eastern record on the Crimean Peninsula (Jaschhof et al. 2011). The current record from Georgia indicates a wider distribution in the Western Palaearctic.

Subfamily Mycetophilinae

Tribe Exechiini

Allodia lugens (Wiedemann, 1817)

Material

4♂♂, SZS-4 (2♂♂ ZFMK, 1♂ IUTG, 1♂ IZBE); 7♂♂, SJ-3; 1♂, SJ-8; 1♂, MM-2; 1♂, MM-11; 1♂, MM-12. Total: 15♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Holarctic.

Remarks

Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Allodia ornaticollis (Meigen, 1818)

Material

1♂, SJ-9; 1♂, MM-12; 1♂, MM-14; 1♂, K-1. Total: 4♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Mtskhetha-Mthianethi, Kakheti.

General distribution

Holarctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 2003).

Allodia truncata Edwards, 1921

Material

1♂, SJ-8. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Holarctic.

Allodiopsis domestica (Meigen, 1830)

Fig. 12A

Material

1♂, A-7; 2♂♂, MM-2; 1♂, MM-8. Total: 4♂♂.

Distribution in Georgia

Adjara, Mtskhetha-Mthianethi.

General distribution

Holarctic.

Remarks

Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Figure 12. 

Habitus of Georgian fungus gnats of the family Mycetophilidae A Allodiopsis domestica (Meigen, 1830) B Rymosia affinis Winnertz, 1863 C Brevicornu griseicolle (Staeger, 1840) D Cordyla fissa Edwards, 1925 E Anatella longisetosa Dziedzicki, 1923 F Mycetophila magnicauda Strobl, 1895 G Epicypta scatophora (Perris, 1849) H Phronia tenuis Winnertz, 1863 I Sceptonia tenuis Edwards, 1925 J Zygomyia humeralis (Wiedemann, 1817) K Dynatosoma reciprocum (Walker, 1848).

Allodiopsis korolevi Zaitzev, 1982

Material

1♂, SZS-4 (ZFMK); 1♂, SJ-9. Total: 2♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Samtskhe-Javakheti.

General distribution

Palaearctic.

Allodiopsis rustica (Edwards, 1941)

Material

2♂♂, SJ-8; 1♂, MM-12. Total: 3♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Anatella longisetosa Dziedzicki, 1923

Fig. 12E

Material

1♂, SZS-3 (ZFMK); 3♂♂, SZS-4 (1♂ ZFMK, 1♂ IUTG, 1♂ IZBE). Total: 4♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Europe.

Anatella metae sp. nov.

Material

See in species description above.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Georgia.

Anatella simpatica Dziedzicki, 1923

Material

1♂, MM-2; 1♂, MM-12. Total: 2♂♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Holarctic.

Brachycampta alternans (Zetterstedt, 1838)

Material

1♂, MM-14. Total: 1♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Holarctic.

Brachycampta czernyi (Landrock, 1912)

Material

1♂, MM-2. Total: 1♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Holarctic.

Brachycampta grata (Meigen, 1830)

Material

4♂♂, I-6; 1♂, I-10; 1♂, SJ-3; 1♂, MM-5; 1♂, MM-8. Total: 8♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 2003).

Brachycampta foliifera (Strobl, 1910)

Material

1♂, MM-2. Total: 1♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Holarctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 2003).

Brachycampta neglecta Edwards, 1925

Material

1♂, I-6. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Palaearctic.

Brachycampta pistillata (Lundström, 1911)

Material

5♂♂, I-6. Total: 5♂♂.

Distribution in Georgia

Imereti.

General distribution

Holarctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 2003).

Brachycampta protenta Laštovka & Matile, 1974

Material

1♂, SZS-4 (ZFMK). Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Holarctic.

Brachycampta westerholti Caspers, 1980

Material

1♂, SJ-8. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Western Palaearctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 2003).

Brevicornu auriculatum (Edwards, 1925)

Material

1♂, SZS-4 (ZFMK); 1♂, A-8. Total: 2♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Adjara.

General distribution

Palaearctic.

Brevicornu bellum (Johannsen, 1912)

Material

1♂, SZS-4 (ZFMK). Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Holarctic.

Brevocornu fuscipenne (Staeger, 1840)

Material

1♂, I-6; 1♂, SJ-8. Total: 2♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti.

General distribution

Holarctic.

Brevicornu griseicolle (Staeger, 1840)

Fig. 12C

Material

8♂♂, SZS-4 (3♂♂ ZFMK, 3♂♂ IUTG, 2♂♂ IZBE); 1♂, I-1; 2♂♂, I-6; 1♂, SK-1; 1♂, SJ-6; 4♂♂, SJ-7; 1♂, SJ-9; 1♂, MM-2; 1♂, MM-14; 1♂, K-1. Total: 21♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Shida Kartli, Samtskhe-Javakheti, Mtskhetha-Mthianethi, Kakheti.

General distribution

Palaearctic.

Brevicornu intermedium (Santos Abreu, 1920)

Material

1♂, I-11, 2♂♂, SJ-7; 2♂♂, SJ-8; 1♂, SJ-9; 1♂, MM-14. Total: 7♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Western Palaearctic.

Brevicornu proximum (Staeger, 1840)

Material

2♂♂, I-6; 1♂, I-11; 1♂, SJ-8; 1♂, SJ-9; 1♂, MM-5; 2♂♂, MM-12; 1♂, MM-13. Total: 9♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 2003).

Brevicornu sericoma (Meigen, 1830)

Material

1♂, SZS-3 (ZFMK); 1♂, SZS-4 (IZBE); 1♂, SJ-7; 4♂♂, SJ-8; 1♂, SJ-9; 3♂♂, KK-1; 1♂, MM-2; 1♂, MM-8; 1♂, MM-14; 1♂, K-2. Total: 15♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Samtskhe-Javakheti, Kvemo Kartli, Mtskhetha-Mthianethi.

General distribution

Holarctic.

Remarks

Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Cordyla brevicornis (Staeger, 1840)

Material

1♂, SZS-3 (IZBE); 5♂♂, SZS-4 (3♂♂ ZFMK, 2♂♂ IUTG); 1♂, A-5; 1♂, A-7; 1♂, I-6; 1♂, SJ-2; 1♂ (IZBE), KK-1; 1♂ 2♀♀, MM-12; 1♂, K-6. Total: 13♂♂ 2♀♀.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Adjara, Imereti, Samtskhe-Javakheti, Kvemo Kartli, Mtskhetha-Mthianethi, Kakheti.

General distribution

Palaearctic.

Cordyla crassicornis Meigen, 1818

Material

1♂, I-1; 1♂, I-2; 1♂, I-6; 1♂, I-11; 1♂, MM-14; 1♂, K-4; 1♂, K-6. Total: 7♂♂.

Distribution in Georgia

Imereti, Mtskhetha-Mthianethi, Kakheti.

General distribution

Palaearctic.

Cordyla fasciata Meigen, 1830

Material

1♂, MM-7. Total: 1♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Cordyla fusca Meigen, 1804

Material

2♂♂, SJ-2 (1♂ ZFMK, 1♂ IZBE). Total: 2♂♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Cordyla fissa Edwards, 1925

Fig. 12D

Material

1♂, I-2; 1♂, I-3 (18.v–1.vi.2013); 1♂, SJ-7; 1♂, KK-1. Total: 4♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti, Kvemo Kartli.

General distribution

Palaearctic.

Cordyla insons Laštovka & Matile, 1974

Material

2♂♂, SJ-2 (1♂ ZFMK, 1♂ IZBE). Total: 2♂♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Cordyla murina (Winnertz, 1863)

Material

2♂♂, I-3 (18.v–1.vi.2013 and 5–19.x.2013); 1♂, SJ-12. Total: 3♂♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Cordyla nitidula Edwards, 1925

Material

1♂, I-14; 1♂, K-5. Total: 2♂♂.

Distribution in Georgia

Imereti, Kakheti.

General distribution

Palaearctic.

Cordyla pusilla Edwards, 1925

Material

3♂♂, SZS-3 (IZBE); 45♂♂, SZS-4 (16♂♂ ZFMK, 16♂♂ IUTG, 13♂♂ IZBE); 2♂♂, I-6; 2♂♂, K-4; 1♂, K-6. Total: 53♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Kakheti.

General distribution

Palaearctic.

Exechia bicincta (Staeger, 1840)

Material

4♂♂, A-5; 3♂♂, A-7; 1♂, I-9; 1♂, I-10; 1♂, I-11; 1♂, I-12; 1♂, MM-8; 1♂, MM-14; 1♂, K-2. Total: 14♂♂.

Distribution in Georgia

Adjara, Imereti, Mtskhetha-Mthianethi, Kakheti.

General distribution

Holarctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 2003).

Exechia dentata Lundström, 1916

Material

1♂, A-7. Total: 1♂.

Distribution in Georgia

Adjara.

General distribution

Europe.

Exechia dorsalis (Staeger, 1840)

Material

1♂, SJ-7; 2♂♂, MM-12; 1♂, MM-13; 3♂♂, MM-14. Total: 7♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Exechia fusca (Meigen, 1804)

Material

2♂♂, I-6; 1♂, SJ-3; 2♂♂, MM-2; 1♂, MM-8; 2♂♂, MM-12. Total: 8♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Holarctic.

Exechia repanda Johannsen, 1912

Material

2♂♂, K-4. Total: 2♂♂.

Distribution in Georgia

Kakheti.

General distribution

Holarctic.

Exechia repandoides Caspers, 1984

Material

1♂, A-3; 1♂, I-3 (24.viii–7.ix.2013); 1♂, SJ-4; 1♂, MM-12; 1♂, K-5. Total: 5♂♂.

Distribution in Georgia

Adjara, Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi, Kakheti.

General distribution

Europe

Exechia seriata (Meigen, 1830)

Material

1♂, A-5. Total: 1♂.

Distribution in Georgia

Adjara.

General distribution

Palaearctic.

Exechiopsis (Exechiopsis) dumitrescae (Burghele-Balacesco, 1972)

Material

1♂, I-12. Total: ♂.

Distribution in Georgia

Imereti.

General distribution

Palaearctic.

Exechiopsis (Exechiopsis) furcata (Lundström, 1911)

Material

1♂ 1♀, MM-1. Total: 1♂ 1♀.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Europe.

Exechiopsis (Exechiopsis) pseudindecisa Laštovka & Matile, 1974

Material

5♂♂, MM-2. Total: 5♂♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Remarks

In Transcaucasia recorded from Armenia (Joost and Plassmann 1985).

Exechiopsis (Exechiopsis) magnicauda (Lundström, 1911)

Material

1♂, MM-8; 2♂♂, MM-11. Total: 3♂♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Europe.

Notolopha cristata (Staeger, 1840)

Material

2♂♂, SJ-8; 1♂, SJ-9. Total: 3♂♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Holarctic.

Pseudexechia tuomikoskii Kjærandsen, 2009

Material

1♂, I-10. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Europe.

Rymosia affinis Winnertz, 1863

Fig. 12B

Material

1♂, SJ-8. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Rymosia fasciata (Meigen, 1804)

Material

1♂, I-9. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Europe.

Stigmatomeria crassicornis (Stannius, 1831)

Material

1♂, A-5; 1♂, SK-1; 1♂, SJ-3; 5♂♂, SJ-8; 1♂, MM-2; 8♂♂, MM-8; 5♂♂, MM-11; 4♂♂, MM-12. Total: 26♂♂.

Distribution in Georgia

Adjara, Shida Kartli, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Holarctic.

Synplasta venosa (Dziedzicki, 1910)

Material

1♂, A-1. Total: 1♂.

Distribution in Georgia

Adjara.

General distribution

Europe.

Tarnania fenestralis (Meigen, 1838)

Material

1♂, SJ-7. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Tribe Mycetophilini

Dynatosoma cochleare Strobl, 1895

Material

1♂, SJ-8. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Dynatosoma fuscicorne (Meigen, 1818)

Material

1♂, SZS-3 (ZFMK). Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Holarctic.

Dynatosoma majus Landrock, 1912

Material

1♀, SJ-8; 4♂♂ 1♀, MM-8; 1♂, K-6. Total: 5♂♂ 2♀♀.

Distribution in Georgia

Samtskhe-Javakheti, Mtskhetha-Mthianethi, Kakheti.

General distribution

Palaearctic.

Dynatosoma nigromaculatum Lundström, 1913

Material

1♀, I-6. Total: 1♀.

Distribution in Georgia

Imereti.

General distribution

Palaearctic.

Dynatosoma reciprocum (Walker, 1848)

Fig. 12K

Material

1♂, SJ-8. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Dynatosoma rufescens (Zetterstedt, 1838)

Material

1♂, K-6. Total: 1♂.

Distribution in Georgia

Kakheti.

General distribution

Europe.

Epicypta limnophila Chandler, 1981

Material

1♂, K-5. Total: 1♂.

Distribution in Georgia

Kakheti.

General distribution

Europe.

Epicypta scatophora (Perris, 1849)

Fig. 12G

Material

1♂, K-4; 7♂♂ 4♀♀, K-5. Total: 8♂♂ 4♀♀.

Distribution in Georgia

Kakheti.

General distribution

Palaearctic.

Epicypta torquata Matile, 1977

Material

1♂, A-1; 1♂ 1♀, I-6; 1♂, MM-14; 1♂; K-2. Total: 4♂♂ 1♀.

Distribution in Georgia

Adjara, Imereti, Mtskhetha-Mthianethi, Kakheti.

General distribution

Western Palaearctic.

Macrobrachius kowarzii Dziedzicki, 1889

Material

1♂, I-6; 1♂, K-4. Total: 2♂♂.

Distribution in Georgia

Imereti, Kakheti.

General distribution

Europe.

Mycetophila adumbrata Mik, 1884

Material

1♂, SJ-8; 1♂, MM-13. Total: 2♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Europe.

Mycetophila alea Laffoon, 1965

Material

1♂, A-3; 35♂♂, I-6; 1♂, I-9; 1♂, I-14; 1♂, SK-1; 1♂, SJ-1 (ZFMK); 3♂♂, SJ-4; 2♂♂, SJ-9; 2♂♂, MM-8; 2♂♂, MM-11; 1♂, KK-1. Total: 50♂♂.

Distribution in Georgia

Adjara, Imereti, Shida Kartli, Samtskhe-Javakheti, Mtskhetha-Mthianethi, Kvemo Kartli.

General distribution

Holarctic.

Mycetophila bialorussica Dziedzicki, 1884

Material

1♂, SZS-4 (ZFMK); 1♂, SJ-4; 1♂, SJ-8; 1♂, SJ-9. Total: 4♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Samtskhe-Javakheti.

General distribution

Palaearctic.

Mycetophila blanda Winnertz, 1863

Material

2♂♂, I-6; 1♂, SJ-9. Total: 3♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti.

General distribution

Palaearctic.

Mycetophila brevitarsata (Laštovka, 1963)

Material

1♂, SZS-4 (ZFMK). Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Palaearctic.

Remarks

Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Mycetophila distigma Meigen, 1830

Material

1♂, SJ-8; 1♂, K-4; Total: 2♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Kakheti.

General distribution

Europe.

Mycetophila edwardsi Lundström, 1913

Material

1♂, SJ-7; 3♂♂, SJ-8; 1♂, MM-8. Total: 5♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Europe.

Mycetophila exstincta Loew, 1869

Material

2♂♂, SZS-3 (1♂ ZFMK, 1♂ IZBE); 1♂, I-14; 2♂♂, K-4. Total: 5♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Kakheti.

General distribution

Holarctic.

Mycetophila formosa Lundström, 1911

Material

1♂, SJ-3; 1♂, SJ-8. Total: 2♂♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Mycetophila fungorum (De Geer, 1776)

Material

6♂♂ 3♀♀, SZS-3 (IZBE); 19♂♂ 10♀♀, SZS-4 (9♂♂ 5♂♀ZFMK, 8♂♂ 5♀♀ IUTG, 2♂♂ IZBE); 1♂, I-3 (29.vi–13.vii.2013); 5♂♂ 2♀♀, I-6; 1♀, I-9; 1♂, I-10; 1♀, SK-1; 2♂♂, SJ-1 (IZBE); 2♀♀, SJ-3; 1♂, SJ-4; 2♂♂ 2♀♀, SJ-7; 4♂♂ 4♀♀, SJ-8; 2♂♂ 4♀♀, SJ-9; 1♂, SJ-10; 2♂♂, MM-3; 1♀, MM-6; 2♂♂ 7♀♀, MM-7; 1♂ 1♀, MM-8; 2♂♂ 2♀♀, MM-11; 1♀, MM-12; 1♂, MM-14; 1♂ 2♀♀, KK-1; 1♂, K-4. Total: 54♂♂ 43♀♀.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Shida Kartli, Samtskhe-Javakheti, Mtskhetha-Mthianethi, Kvemo Kartli, Kakheti.

General distribution

Holarctic (extending to the Oriental region).

Mycetophila confluens Dziedzicki, 1884

Material

4♂♂, SJ-8; 1♂, MM-2. Total: 5♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Holarctic.

Mycetophila curviseta Lundström, 1911

Material

4♂♂, SZS-4 (2♂♂ ZFMK, 1♂ IUTG, 1♂ IZBE); 1♂, I-3 (1–15.vi.2013); 3♂♂, I-6; 2♂♂, I-9; 1♂, I-12; 2♂♂, SK-1; 6♂♂, SJ-8; 5♂♂, SJ-9; 1♂, KK-1; 3♂♂, MM-8; 1♂, MM-13; 7♂♂, MM-14; 3♂♂, K-4; 1♂, K-5. Total: 40♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Shida Kartli, Samtskhe-Javakheti, Kvemo Kartli, Mtskhetha-Mthianethi, Kakheti.

General distribution

Palaearctic.

Mycetophila deflexa Chandler, 2001

Material

2♂♂, SZS-3 (1♂ ZFMK, 1♂ IZBE); 1♂, SZS-4 (IUTG). Total: 3♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Europe.

Mycetophila dentata Lundström, 1915

Material

1♂, K-4. Total: 1♂.

Distribution in Georgia

Kakheti.

General distribution

Holarctic.

Mycetophila gentilicia Zaitzev, 1999

Material

1♂, SJ-8. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Mycetophila gibbula Edwards, 1925

Material

1♂, SJ-3. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Mycetophila hetschkoi Landrock 1918

Material

1♂, SZS-4 (ZFMK); 1♂, MM-11; 3♂♂, MM-14. Total: 5♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Mycetophila hyrcania Laštovka & Matile, 1969

Material

1♂, SZS-3 (ZFMK); 1♂, SJ-6; 1♂, KK-1; 3♂♂, K-4. Total: 6♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Samtskhe-Javakheti, Kvemo Kartli, Kakheti.

General distribution

Western Palaearctic.

Mycetophila ichneumonea Say, 1823

Georgian source

Jürgenstein et al. 2015: 30.

Material

6♂♂, SZS-4 (3♂♂ ZFMK, 2♂♂ IUTG, 1♂ IZBE); 1♂, SK-1; 1♂, SJ-2 (IZBE); 1♂, SJ-5; 3♂♂, SJ-8; 1♂, SJ-9. Total: 13♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Shida Kartli, Samtskhe-Javakheti.

General distribution

Holarctic. Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Mycetophila idonea Laštovka, 1972

Georgian source

Jürgenstein et al. 2015: 31–32.

Material

5♂♂, SZS-3 (2♂♂ ZFMK, 2♂♂ IUTG, 1♂ IZBE); 2♂♂, A-1; 1♂, A-3; 1♂, A-7; 10♂♂, I-6; 1♂, I-10; 1♂, SK-1; 1♂, SJ-1 (IZBE); 1♂, KK-1; 1♂, MM-7; 1♂, MM-8; 4♂♂, MM-11; 1♂, MM-14; 1♂, K-4; 1♂, K-5; 1♂, K-6. Total: 33♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Adjara, Imereti, Shida Kartli, Samtskhe-Javakheti, Kvemo Kartli, Mtskhetha-Mthianethi, Kakheti.

General distribution

Europe.

Mycetophila lamellata Lundström, 1911

Material

4♂♂, SJ-7; 1♂, SJ-8; 2♂♂, MM-14. Total: 7♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Europe.

Mycetophila lastovkai Caspers, 1984

Material

1♂, SZS-4 (ZFMK). Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Europe.

Mycetophila luctuosa Meigen, 1830

Material

1♂, SZS-4 (ZFMK); 1♂, I-10; 1♂, MM-4. Total: 3♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Mtskhetha-Mthianethi.

General distribution

Holarctic.

Mycetophila lunata Meigen, 1804

Material

1♂, A-1; 1♂, MM-12. Total: 2♂♂.

Distribution in Georgia

Adjara, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Mycetophila magnicauda Strobl, 1895

Fig. 12F

Material

1♂, SJ-4; 4♂♂, SJ-8; 1♂, SJ-9. Total: 6♂♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Mycetophila marginata Winnertz, 1863

Material

1♂, I-6; 1♂, I-10; 4♂♂, SJ-7; 5♂♂, SJ-8; 2♂♂, SJ-9; 2♂♂, MM-2. Total: 15♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Europe.

Mycetophila morosa Winnertz, 1863

Material

1♂, MM-14. Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Holarctic.

Mycetophila nigrofusca Dziedzicki, 1884

Material

1♂, I-5; 1♂, MM-14. Total: 2♂♂.

Distribution in Georgia

Imereti, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Mycetophila ocellus Walker, 1848

Material

1♂, I-10; 3♂♂, SJ-8; 2♂♂, MM-12. Total: 6♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Holarctic.

Mycetophila occultans Lundström, 1913

Material

6♂♂, SZS-3 (2♂♂ ZFMK, 2♂♂ IUTG, 2♂♂ IZBE);1♂, I-6; 1♂, SJ-8; 1♂, SJ-9; 57♂♂, MM-13; 1♂, MM-14. Total: 67♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Europe.

Mycetophila ornata Stephens, 1829

Material

2♂♂, SJ-8; 1♂, SJ-9; 2♂♂, MM-12. Total: 5♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Remarks

In Transcaucasia recorded from Azerbaijan (Zaitzev 2003).

Mycetophila pictula Meigen, 1830

Material

2♂♂, SJ-9. Total: 2♂♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Holarctic.

Mycetophila pumila Winnertz, 1863

Material

1♂, SZS-4 (ZFMK); 2♂♂, A-1; 2♂♂, A-3; 3♂♂, I-6; 1♂, I-9; 1♂, I-10; 1♂, MM-11; 4♂♂, MM-14; 1♂, K-2. Total: 16♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Adjara, Imereti, Mtskhetha-Mthianethi, Kakheti.

General distribution

Palaearctic.

Mycetophila pseudoforcipata Zaitzev, 1998

Material

1♂, SJ-8; 1♂, SJ-9. Total: 2♂♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Mycetophila ruficollis Meigen, 1818

Georgian source

Jürgenstein et al. 2015: 33.

Material

1♂, K-6. Total: 1♂.

Distribution in Georgia

Kakheti.

General distribution

Palaearctic.

Mycetophila scotica Edwards, 1941

Material

1♂, I-12. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Holarctic.

Mycetophila sigillata Dziedzicki, 1884

Material

3♂♂, I-6; 2♂♂, SJ-4. Total: 5♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti.

General distribution

Holarctic.

Remarks

Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Mycetophila sigmoides Loew, 1869

Material

1♂, I-6. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Holarctic.

Mycetophila signata Meigen, 1830

Material

10♂♂, I-6; 1♂, I-14; 1♂, SJ-4. Total: 12♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti.

General distribution

Palaearctic.

Mycetophila signatoides Dziedzicki, 1884

Material

1♂, SZS-4 (ZFMK); 2♂♂, I-6. Total: 3♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti.

General distribution

Western Palaearctic (see also comment in Kjaerandsen et al. 2007).

Mycetophila sordida van der Wulp, 1874

Material

1♂, SZS-3 (ZFMK); 3♂♂, SZS-4 (1♂ ZFMK, 1♂ IUTG, 1♂ IZBE); 2♂♂, I-6; 1♂, SJ-4; 4♂♂, SJ-8; 1♂, SJ-9; 6♂♂, KK-1; 1♂, MM-11. Total: 19♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Samtskhe-Javakheti, Kvemo Kartli, Mtskhetha-Mthianethi.

General distribution

Holarctic.

Mycetophila strigatoides Landrock, 1927

Material

4♂♂, SZS-3 (2♂♂ ZFMK, 2♂♂ IZBE); 42♂♂, SZS-4 (13♂♂ ZFMK, 16♂♂ IUTG, 13♂♂ IZBE); 1♂, I-10; 1♂, SJ-1 (IZBE); 1♂, SJ-2 (ZFMK). Total: 49♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Samtskhe-Javakheti.

General distribution

Palaearctic.

Mycetophila stylata (Dziedzicki, 1884)

Material

9♂♂, SJ-8; 4♂♂, SJ-9. Total: 13♂♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Mycetophila sublunata Zaitzev, 1998

Material

1♂, SJ-9. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Europe.

Mycetophila subsigillata Zaitzev, 1999

Material

1♂, SJ-8. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Mycetophila sumavica (Laštovka, 1963)

Material

1♂, I-10. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Europe.

Mycetophila trinotata Staeger, 1840

Material

1♂, SZS-3 (ZFMK); 5♂♂, MM-13; 12♂♂, MM-14; 1♂, K-2. Total: 19♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Mtskhetha-Mthianethi, Kakheti.

General distribution

Holarctic.

Mycetophila uliginosa Chandler, 1988

Material

1♂, SZS-4 (ZFMK). Total: 1♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Europe.

Mycetophila unicolor Stannius, 1831

Material

3♂♂, A-1; 2♂♂, A-7; 1♂, SJ-2 (ZFMK). Total: 6♂♂.

Distribution in Georgia

Adjara, Samtskhe-Javakheti.

General distribution

Western Palaearctic.

Phronia basalis Winnertz, 1863

Material

1♂, MM-10. Total: 1♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Western Palaearctic.

Phronia biarcuata (Becker, 1908)

Material

1♂, SJ-8; 1♂, SJ-9; 1♂, MM-12. Total: 3♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Holarctic.

Remarks

In Transcaucasia recorded from Armenia (Joost and Plassmann 1985).

Phronia conformis (Walker, 1856)

Material

2♂♂, I-6. Total: 2♂♂.

Distribution in Georgia

Imereti.

General distribution

Holarctic.

Phronia electa Dziedzicki, 1889

Material

1♂, SJ-8. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Phronia exigua (Zetterstedt, 1852)

Material

2♂♂, MM-2; 1♂, MM-8. Total: 3♂♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Holarctic.

Remarks

Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Phronia humeralis Winnertz, 1863

Material

1♂, A-7; 1♂, SJ-4; 2♂♂, SJ-8; 2♂♂, SJ-9. Total: 6♂♂.

Distribution in Georgia

Adjara, Samtskhe-Javakheti.

General distribution

Palaearctic.

Phronia forcipata Winnertz, 1863

Material

1♂, SZS-4 (ZFMK); 8♂♂, I-6. Total: 9♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti.

General distribution

Palaearctic.

Remarks

Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Phronia nitidiventris (van der Wulp, 1859)

Material

2♂♂, I-6; 1♂, I-9; 1♂, I-10. Total: 4♂♂.

Distribution in Georgia

Imereti.

General distribution

Palaearctic.

Remarks

Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Phronia notata Dziedzicki, 1889

Material

1♂, SJ-8. Total: 1♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Palaearctic.

Phronia obtusa Winnertz, 1863

Material

1♂, I-6. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Holarctic.

Phronia petulans Dziedzicki, 1889

Material

6♂♂, MM-12. Total: 6♂♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Holarctic.

Phronia signata Winnertz, 1863

Material

9♂♂, I-6; 1♂, I-9; 4♂♂, I-10; 1♂, SJ-9; 8♂♂, MM-8; 1♂, MM-9. Total: 24♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Remarks

Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Phronia tenuis Winnertz, 1863

Fig. 12H

Material

2♂♂, SZS-4 (1♂ ZFMK, 1♂ IZBE); 1♂, I-10; 1♂, MM-12. Total: 4♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Mtskhetha-Mthianethi.

General distribution

Holarctic, extending to the Oriental region.

Phronia triangularis Winnertz, 1863

Material

1♂, I-6; 1♂, SJ-8; 1♂, MM-2; 1♂, MM-8. Total: 4♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Western Europe.

Platurocypta testata (Edwards, 1925)

Material

1♂, I-3 (13–27.vi.2013); 2♂♂, SJ-4; 1♂, K-4. Total: 4♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti, Kakheti.

General distribution

Holarctic.

Platurocypta punctum (Stannius, 1831)

Material

1♂, K-2. Total: 1♂.

Distribution in Georgia

Kakheti.

General distribution

Holarctic.

Sceptonia cryptocauda Chandler, 1991

Material

18♂♂, MM-13; 18♂♂, MM-14; 8♂♂, K-2. Total: 44♂♂.

Distribution in Georgia

Mtskhetha-Mthianethi; Kakheti.

General distribution

Western Palaearctic.

Sceptonia demeijerei Bechev, 1997

Material

1♂, MM-5. Total: 1♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Europe.

Sceptonia flavipuncta Edwards, 1925

Material

1♂, SZS-3 (IZBE); 6♂♂, I-6; 3♂♂, I-9; 2♂♂, I-14; 1♂, SJ-2 (ZFMK); 1♂, KK-1; 10♂♂, MM-13; 14♂♂, MM-14. Total: 38♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Samtskhe-Javakheti, Kvemo Kartli, Mtskhetha-Mthianethi.

General distribution

Europe.

Sceptonia humerella Edwards, 1925

Material

1♂, SJ-3; 1♂, SJ-4. Total: 2♂♂.

Distribution in Georgia

Samtskhe-Javakheti.

General distribution

Europe.

Sceptonia membranacea Edwards, 1925

Material

5♂♂, MM-13. Total: 5♂♂.

Distribution in Georgia

Mtskhetha-Mthianethi.

General distribution

Europe.

Sceptonia nigra (Meigen, 1804)

Material

2♂♂, A-3; 1♂, MM-13; 14♂♂, MM-14; 7♂♂, K-2. Total: 24♂♂.

Distribution in Georgia

Adjara, Mtskhetha-Mthianethi, Kakheti.

General distribution

Palaearctic.

Sceptonia tenuis Edwards, 1925

Fig. 12 I

Material

1♂, SK-1; 3♂♂, SJ-2 (1♂ ZFMK, 1♂ IUTG, 1♂ IZBE); 2♂♂, MM-12. Total: 6♂♂.

Distribution in Georgia

Shida Kartli, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Europe.

Trichonta aberrans Lundström, 1911

Material

1♂, I-6. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Europe.

Trichonta atricauda (Zetterstedt, 1852)

Material

1♂, I-6. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Holarctic.

Trichonta clavigera Lundström, 1913

Material

2♂♂, I-6; 1♂, KK-1; 1♂, MM-12. Total: 4♂♂.

Distribution in Georgia

Imereti, Kvemo Kartli, Mtskhetha-Mthianethi.

General distribution

Palaearctic.

Trichonta falcata Lundström, 1911

Material

2♂♂, A-7. Total: 2♂♂.

Distribution in Georgia

Adjara.

General distribution

Holarctic.

Remarks

Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Trichonta fragilis Gagne, 1981

Material

1♂, I-6. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Holarctic.

Trichonta perspicua van der Wulp, 1881

Material

1♂, I-17. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Holarctic.

Trichonta subterminalis Zaitzev & Menzel, 1996

Material

4♂♂, I-6; 1♂, SK-1. Total: 5♂♂.

Distribution in Georgia

Imereti, Shida Kartli.

General distribution

Palaearctic.

Trichonta trifida Lundström, 1909

Material

1♂, I-10. Total: 1♂.

Distribution in Georgia

Imereti.

General distribution

Northern Europe.

Remarks

Kjæranden and Søli (2020) recently reinstated the species and provided detailed figures of the male terminalia of the allied species.

Trichonta vitta (Meigen, 1830)

Material

9♂♂, I-6; 1♂, SJ-4; 1♂, SJ-8; 2♂♂, K-4; 2♂♂, K-5. Total: 15♂♂.

Distribution in Georgia

Imereti, Samtskhe-Javakheti, Kakheti.

General distribution

Holarctic.

Trichonta vulgaris Loew, 1869

Material

1♂, SZS-4 (ZFMK); 2♂♂, SJ-8; 1♂, SJ-9. Total: 4♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Samtskhe-Javakheti.

General distribution

Holarctic.

Remarks

Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Zygomyia humeralis (Wiedemann, 1817)

Fig. 12J

Material

1♂, SZS-3 (IZBE); 5♂♂, SZS-4 (2♂♂ ZFMK, 2♂♂ IUTG, 1♂ IZBE); 1♂, SJ-4; 2♂♂, MM-12; 1♂, MM-13; 3♂♂, MM-14. Total: 13♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Samtskhe-Javakheti, Mtskhetha-Mthianethi.

General distribution

Europe.

Zygomyia pseudohumeralis Caspers, 1980

Material

1♂, SJ-2 (ZFMK); 1♂, SJ-4; 1♂, SJ-7; 1♂, KK-1; 5♂♂, K-4. Total: 9♂♂.

Distribution in Georgia

Samtskhe-Javakheti, Kvemo Kartli, Kakheti.

General distribution

Palaearctic.

Zygomyia semifusca (Meigen, 1818)

Material

1♂, SZS-4 (ZFMK); 1♂, I-6; 1♂, SJ-3; 1♂, SJ-5. Total: 4♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Imereti, Samtskhe-Javakheti.

General distribution

Holarctic.

Zygomyia setosa Barendrecht, 1938

Material

1♂, K-4. Total: 1♂.

Distribution in Georgia

Kakheti.

General distribution

Europe.

Remarks

A very rare species with a scattered distribution in Europe: recorded from the Netherlands, Germany and Switzerland (Chandler 2013). There is an unpublished record also from Estonia (personal observation).

Zygomyia valida Winnertz 1863

Material

10♂♂, SZS-4 (4♂♂ ZFMK, 3♂♂ IUTG, 3♂♂ IZBE). Total: 10♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi.

General distribution

Palaearctic.

Zygomyia vara (Staeger, 1840)

Material

1♂, SZS-2; 1♂, SZS-4 (ZFMK); 1♂, A-1; 2♂♂, SJ-9. Total: 5♂♂.

Distribution in Georgia

Samegrelo-Zemo Svanethi, Adjara, Samtskhe-Javakheti.

General distribution

Holarctic.

Remarks

Listed to occur in Transcaucasia without further details (Zaitzev 2003).

Discussion

This is the first attempt to provide a synoptic list of Sciaroidea species of the Transcaucasian region. However, the recorded 246 species (245 from original study + one from literature data) of fungus gnats are the result of a preliminary survey, while further sweeping studies will probably increase that number considerably (see also below). As expected, the majority of the recorded species are widely distributed in the Palaearctic or Holarctic regions (38% and 26% of the recorded species, respectively), while 22% of species are restricted to Europe and 7% to the Western Palaearctic (Fig. 13). In addition, one species was so far known only from the Eastern Palaearctic and 17 species (11 described and 6 undescribed) are classified (tentatively) as Caucasian endemics. These proportions can change as fungus gnats are rather poorly known in several Palaearctic regions including the East Palaearctic, Asia Minor, Central Asia, as well as other regions in Caucasia.

Figure 13. 

Grouping of the recorded Georgian fungus gnat species in accordance with their known distribution.

The estimated species richness is the highest when calculated using Jackknife-2 nonparametric estimator (404 species, Fig. 14). This method has been discussed as possibly overestimating the true richness (e.g. Poulin 1998). On the other hand, Smith and van Belle (1984) showed that both Jackknife and Bootstrap estimators underestimate the actual number of species if there is a large number of rare species considered and number of samples is low. That can also be the case in the current data as the number of recorded singletons and doubletons is exceptionally high (82 and 42 species, respectively) and the number of species recorded from one sample only (= unique species) constitutes 43% of the observed diversity (107 out of the 251). Within the listed species, only nine were recorded from more than ten samples and 31 species from 5–10 samples. To compare, relatively well studied countries of similar size in Central and Northern Europe (e.g. Czech Republic, Slovak Republic, Estonia) have roughly 600 fungus gnat species recorded (Ševčík and Košel 2009, Ševčík and Kurina 2011a, b, pers. observation). Taking into account the mountainous landscape, high diversity of habitats, microclimates in Georgia and that several regions were not covered by sampling of the current study (see Fig. 1), it can be presumed that the observed 245 species (+ one based on the literature data) do not constitute more than half of the actual diversity, probably less.

Figure 14. 

Species accumulation curves (EstimateS, Vesrion 9.1.0.). Three nonparametric estimators (Chao 2, Jackknife 2 and Bootstrap) of total species richness are calculated. S(est) is the cumulative number of species observed.

Surprisingly, the most abundant species was Synapha fasciata (769 specimens from 19 samples) followed by Orfelia georgica (175 specimens from 14 samples). In the European boreal and temperate regions, the most abundant species belong frequently to the subfamily Mycetophilinae and/or to the genera Boletina Staeger and Mycomya Rondani. Several of the recorded species considerably increase the knowledge of their distribution, the most remarkable of them include: Neoempheria brevilineata (earlier from Japan only), Clastobasis loici (earlier from Europe and Japan), Lusitanoneura chandleri (earlier from the Mediterranean Islands), Zygomyia setosa (earlier with scattered distribution in Europe), Manota unifurcata (earlier from Europe only).

From the material underlining this study, four new species have been described earlier (Kurina and Jürgenstein 2013; Kurina 2018; Ševčík et al. 2020), three new species are described above and six putatively new species are left to be described in the future due to insufficiency of the available material or its quality. More exhaustive sampling will naturally yield a number of new taxa to be described in the future.

Acknowledgements

I am grateful to Prof. T. Tammaru (University of Tartu, Estonia) for organizing the collecting trips to Georgia in 2012 and 2013, and to Prof. G. Japoshvili (Agricultural University of Georgia, Tbilisi, Georgia) for help during the fieldwork including arrangement of permits. The study visit to the Zoological Research Museum Alexander Koenig, Bonn, Germany was funded by the European Commission’s Research Infrastructure programme SYNTHESYS+ (DE-TAF-2498). I am grateful to X. Mengual (Bonn, Germany), U. Jürivete (Tallinn, Estonia) and A. Selin (Tallinn, Estonia) for providing specimens for this study. J. Kjærandsen (Tromsø, Norway), J. Salmela (Rovaniemi, Finland) and A. Polevoi (Petrozavodsk, Russia) are thanked for comments on particular species. I am much obliged to S. Fitzgerald (Corvallis, U.S.A.) for linguistic suggestions on the manuscript. A. Polevoi (Petrozavodsk, Russia), Jan Ševčík (Ostrava, Czech Republic) and P. Chandler (Melksham, United Kingdom) suggested valuable improvements to the manuscript.

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