Butterflies (Lepidoptera: Papilionoidea) of Georgia (Caucasus): annotated review of regional butterfly fauna with vernacular names index, notes on distribution and phenology

Iwona Słowińska; Krzysztof Jonko

doi:10.7717/peerj.18720

Butterflies (Lepidoptera: Papilionoidea) of Georgia (Caucasus): annotated review of regional butterfly fauna with vernacular names index, notes on distribution and phenology

1Department of Invertebrate Zoology and Hydrobiology, Faculty of Biology and Environmental Protection, University of Lodz, Łódź, Poland

2Łódź, Poland

DOI: 10.7717/peerj.18720

Published: 2025-02-17
Accepted: 2024-11-26
Received: 2024-05-09

Academic Editor: Daniel Silva

Subject Areas: Biodiversity, Ecology, Entomology, Taxonomy, Zoology
Keywords: Caucasus, Georgia, Biodiversity, Butterflies, Checklist, Vernacular names, Regional distribution

Copyright: © 2025 Słowińska and Jonko
Licence: This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, reproduction and adaptation in any medium and for any purpose provided that it is properly attributed. For attribution, the original author(s), title, publication source (PeerJ) and either DOI or URL of the article must be cited.

Cite this article: Słowińska I, Jonko K. 2025. Butterflies (Lepidoptera: Papilionoidea) of Georgia (Caucasus): annotated review of regional butterfly fauna with vernacular names index, notes on distribution and phenology. PeerJ 13:e18720 https://doi.org/10.7717/peerj.18720

The authors have chosen to make the review history of this article public.

Abstract

Background

It is well known that butterflies are valuable indicators of environmental quality, given their existence in various habitats. Collecting and regularly updating data on species richness, abundance, and distribution of all butterfly species in the country is crucial for effective monitoring and conservation efforts, which can ultimately help minimise biodiversity losses. Since the last publication of the Georgian butterfly list, there have been numerous reports registering taxonomic revisions, nomenclatural changes or providing several new butterfly “cryptic species” based on genetic research. In the following article, based on a review of various sources of data, including existing literature and new, unpublished data, we present an annotated regional checklist of butterflies of Georgia, a country that is a part of the Caucasus ecoregion representing one of 36 biodiversity “hotspots”.

Methods

A database with all reported species for Georgia was created by compiling information from critical reviews of all available literature reports, records submitted by contributors of three websites dedicated to butterfly fauna, and data deposited in the Global Biodiversity Information Facility (GBIF) database. Various specialised sources were used to extract vernacular species names (Georgian, Russian and English).

Results

The updated list of butterflies from Georgia includes 244 species of the superfamily Papilionoidea recorded from almost 600 different locations. Nearly 25% of the 244 species were considered rare and extremely rare, while at the same time being at risk of potential extinction. For each species, we present brief phenological information, distribution in Georgia/Caucasus, occurrence status in each region of the country, thumbnails (ventral and dorsal view), as well as a list of vernacular names in Georgian, Russian and English. Regarding species that are reported in the literature as new, uncertain, or questionable in Georgia, we provide the relevant comments. In comparison to the other republics of Transcaucasia (Armenia, Azerbaijan), we noted a similar number of species. Our studies provide a robust baseline of data for further exploration of the Lepidoptera fauna of Georgia. This foundation should help to fill in the gaps in knowledge regarding regional species distribution, phenology, and habitat requirements.

Introduction

Insect abundance and diversity have declined dramatically during the last decades (Hallmann et al., 2017, 2021; Leather, 2017; Forister, Pelton & Black, 2019; Rhodes, 2019; Sánchez-Bayo & Wyckhuys, 2019; Dalton et al., 2023). The conservation of biodiversity has emerged as a critical global concern. Data on species distribution and fauna diversity may serve as points of reference for estimating environmental changes and, at the same time, constitute the basis for assessing the conservation status of species and implementing conservation strategies to prevent further loss of insects, including Lepidoptera. Regarding Georgian fauna, Didmanidze (2004) is the only work known to the authors with a clear focus to document the lepidopteran fauna of Georgia. It was the first checklist of Georgian butterflies containing all species found in this country known at that time. Unfortunately, many species listed by Didmanidze have been considered doubtful or misidentified by, among others, Nekrutenko, Korshunov & Effendi (1982a, 1982b) and Tshikolovets & Nekrutenko (2012) and require independent confirmation. Generally, data on Georgian butterflies are still quite scattered and mostly deal with some separate regions and taxa only.

The Republic of Georgia is situated between two major mountain ranges: the Greater Caucasus and the Lesser Caucasus. Along with Azerbaijan and Armenia, Georgia forms one Transcaucasian region between the Caucasus Mountains and the borders of Iran and Turkey. It is divided into two autonomous republics (Abkhazia, Adjara), one city district (Tbilisi) and nine regions (Fig. 1).

Figure 1: Map of Georgia with administrative divisions.
Map credit: Krzysztof Jonko.

Download full-size image

DOI: 10.7717/peerj.18720/fig-1

Georgia is located in the Caucasus ecoregion—one of 36 biodiversity hot-spots—which shelters one of the most diverse fauna of the temperate region (Zazanashvili et al., 2004). This ecoregion is distinguished for having high levels of diversity and endemism while, at the same time, being seriously threatened by loss of habitats. Although Georgia covers a small part of the Caucasus, it contains the largest part of Caucasian forests, which cover about 40% of its territory. As many as 98% of Georgian forests are growing in the mountains, and only 2% is situated in low-lying areas (Patarkalashvili, 2017). The major threats to these habitats include overcutting, overgrazing, forest pests and diseases, fires, invasive alien species, and poor land-use practices (Akhalkatsi, 2015; Zazanashvili et al., 2020; Beridze & Dering, 2021). These factors, along with increasing climate changes intensify the overall loss of habitats and, in consequence, the biodiversity in Georgia. For biodiversity management, a national checklist of insect species is essential. Such initiatives as regularly updatable checklists or databases have already been compiled for many countries. These projects may be an important source of necessary information for a broad range of potential users, including conservationists, citizen scientists and scientists that would like to understand patterns of insect species diversity and distribution at a national or regional level.

The history of butterfly research in the region known as present-day Georgia starts in 1832 with a article by Ménétriés (1832) who provided the first species list of the Caucasus butterflies, though without precise locality data. For this reason, that publication could not be used as a data source for the regional checklist of butterflies of Georgia. Some time later, Nordmann (1851) published the first list of Georgian Lepidoptera species with full locality data. More intensive studies of butterflies from the country began in the second half of the 19^th century (Lederer, 1864, 1870; Hedemann, 1876; Christoph, 1877, 1881, 1886, 1889; Romanoff, 1884, 1885, 1887; Shavrov, 1886; Radde, 1899) and continued quite actively in the first half of the 20^th century. The period before the end of the Second World War was characterised by many reports of Georgian butterflies (e.g., Jachontov, 1911; Sheljuzhko, 1924; Miljanovskiy, 1941, 1947), but none have been so extensive and comprehensive as publications by Romanoff (1884, 1885, 1887). After the Second World War, many different authors, mainly from the former Soviet Union, published data on butterflies from the territory of Georgia (e.g., Rjabov, 1958; Miljanovskiy, 1964; Batiashvili & Didmanidze, 1973; Didmanidze, 1975, 1979, 1980; Vashakidze, 1976; Nekrutenko, 1990). Studies on Georgian butterflies diminished in the late 20^th century and have only recently resumed (e.g., Didmanidze et al., 2002; Tshikolovets, 2003, 2011; Didmanidze, 2004, 2005a, 2005b; Korb, 2005; Morgun, 2010, 2019; Tshikolovets & Nekrutenko, 2012; Bolshakov & Korb, 2013; Stradomsky & Tikhonov, 2015; Korb & Bolshakov, 2016). Currently, lepidopterology in Georgia faces significant challenges due to the absence of successors to Professor Didmanidze in both scientific institutions and among actively involved enthusiasts. Mumladze, Japoshvili & Anderson (2019) indicate that only a few taxa—such as Coleoptera, Hemiptera, Hymenoptera, Acari (Oribatida), and Gastropoda—are actively studied in Georgia. Furthermore, there is a lack of comprehensive data regarding the distribution of taxa, particularly butterflies.

Twenty years have passed since Didmanidze’s checklist was published (Didmanidze, 2004); therefore, an update is needed, especially since several new investigations have been carried out, recording new species or new localities for species previously noted in Georgia. Intense genetic studies in recent years have resulted in changes in systematics and increased the number of reported species not previously listed. Currently, apart from the information from Didmanidze (2004), Tshikolovets & Nekrutenko (2012) and scarce data for individual species from four websites, detailed data on the distribution of most butterfly species in Georgia are not available. Based on the number of recorded taxa, several other species can be expected to occur in Georgia, including those reported in surrounding countries or new to science.

Therefore, in the present article, we provide an updated and annotated regional list of Georgian butterflies with vernacular names, species distribution in Georgia/Caucasus, occurrence status in each region of the country, phenological data and habitat requirements. Moreover, we present additional comments for the species with doubtful presence in Georgia and for the species that have not been previously reported in Georgia. This paper should assist in the understanding of present-day knowledge of Georgia’s butterfly fauna and form a starting point for further studies into the biodiversity and conservation of butterflies in that country.

Materials and Methods

Literature records

We need to point out that the checklist by Didmanidze (2004) contains errors such as redundant or missing letters in Latin names, nomenclatural inaccuracies, mistaken authors of species names, omissions, and incomplete or incorrect information about ranges of species. For that reason, we used Tshikolovets & Nekrutenko (2012) as our reference data source. We considered that the aforementioned authors summarised the important publications on the butterfly fauna of the Caucasus. Nevertheless, we also critically reviewed all available literature reports to verify original spelling, possible taxonomic changes and precise distributional information in each region (e.g., Eversmann, 1851; Alphéraky, 1876; Staudinger & Rebel, 1901; Moucha, 1968; Batiashvili & Didmanidze, 1973; Gross, 1978; Tuzov, 1993; Tuzov et al., 1997, 2000; Bukhnikashvili et al., 2012). We did not verify material for misidentified or doubtful species, as it was beyond the scope of the present study. Nonetheless, we have decided to include those records in the checklist. All species with uncertain or questionable occurrence in Georgia are marked with an asterisk (*) placed at the species name in all presented tables. Nevertheless, we included some comments about those species, as well as about some new findings. Due to the lack of data on the number of individuals of particular species, especially in older publications, we defined species rarity using a criterion that focuses on their frequency of occurrence within the territory of Georgia. If a species was recorded from no more than three localities, we considered it rare; if it was recorded from only one locality, we regarded it as extremely rare within the Georgian fauna.

It is important to note that lepidopteran taxonomy and nomenclature are active fields of study, with ongoing revisions based on new discoveries and genetic analyses. The classification system continues to evolve, and for that reason we decided to base our study on the classification and nomenclature used in the recent publication by Rajaei et al. (2023). The primary source of Russian vernacular names was the publication by Lvovsky & Morgun (2007), while Georgian names were derived from various sources, primarily Georgian publications (including Bukhnikashvili et al., 2012) and the portal https://biodiversity.iliauni.edu.ge./.

GBIF records

The records obtained from the Global Biodiversity Information Facility (https://www.gbif.org/, accessed on 2 April 2022) have been subjected to a very critical assessment. All records containing obvious errors have been excluded—for example, records collected from New Georgia Island or from the state of Georgia (in the United States of America) which were erroneously assigned (in the GBIF) to Georgia in the Caucasus. The detailed locations of all GBIF records were also verified. All records with missing Global Positioning System (GPS) coordinates or those indicating localities outside the border of present-day Georgia (i.e., localities in the territory of Russia or Turkey) have been excluded. All records described as human observation that did not contain an image confirming such observation were also excluded. All human observations, confirmed by photographs of species that could be confused with other species, were additionally verified.

Other sources of records

We had access to records submitted by contributors to the Lepidoptera Mundi website (Jonko & Słowińska, 2023) and two other websites: the Georgian Biodiversity Database (Tarkhnishvili & Chaladze, 2023), Butterflies of the Caucasus and the South of Russia (Tikhonov et al., 2023). Additionally, we used data based on the material photographed by us during the Caucasus expeditions in the years 2018–2019 under the project “V4 & Eastern Partnership”. All observed individuals were photographed and identified in the field, and then they were included in the general database.

Database and visualisation

For the purpose of our study, a Microsoft Access 2010 relational database was created locally on the authors’ computers and subsequently migrated to the MySQL platform, based on all available data sources. The database consists of several tables containing information about taxonomy, regions, locations, vernacular names, dates of individual records and the period in which the record was noted originally. It includes taxa identified at least to the species level and records with detailed localities or at least region name. The final checklist has been created for the species level. All taxa within families are arranged in systematic order. The database of faunistic records in SQL/MS Access format is available from the authors upon request.

Visualisation of the regional distribution maps of all species was generated using the GD library and scripts written in PHP. Moreover, the thumbnails of dorsal and ventral views of each species are presented (where available). The main source of the thumbnails is the authors’ website Lepidoptera Mundi (Jonko & Słowińska, 2023). All of the thumbnails were created from photos of pinned specimens, with the photos edited in Adobe Photoshop CS6. We note that the presented thumbnails are not all generated to the same scale and do not contain a 1 cm reference bar. These images are used only for the depiction of a species and may not represent individuals collected in the Caucasus.

To clearly separate historical and more recent data, we decided to divide all records in the database into three periods: very old (historical) records up to 1945 (end of the Second World War), records collected between the end of the Second World War and Georgia’s independence (1945–1991), and records collected since Georgia gained its independence (post-1991). This division was made not for any scientific reasons, but solely based on historical events.

Results

List of Georgian butterflies

Based on the most up-to-date butterfly checklist of the Caucasus by Tshikolovets & Nekrutenko (2012), we registered 201 species confirmed in Georgia. The current systematics forced a recalculation of the number of species listed by the aforementioned authors to make it comparable with the current state of knowledge. In summary, prior to the present study, 205 species of butterflies representing five families were recorded in Georgia. In the present article, 39 species were added to this list and, as a result, a total of 244 species belonging to 85 genera are listed for Georgia. The greatest species richness was recorded within the families Nymphalidae (96 species, 34 genera) and Lycaenidae (83 species, 26 genera) (Table 1).

Table 1:

Summary of the number of subfamilies, genera and species within each butterfly family in Georgia.

Family	Subfamily	Genera	Species	Endemic*
Hesperiidae	3	12	29	1
Lycaenidae	3	26	83	6
Nymphalidae	6	34	96	7
Papilionidae	2	4	8	1
Pieridae	3	9	28	1
Total	17	85	244	16

DOI: 10.7717/peerj.18720/table-1

Note:

* Endemic to the Caucasus.

Sixteen species (6.6% of Georgian fauna) were considered Caucasian endemics (see Tables 1 and 2), while four species (Carterocephalus palaemon, Parnassius phoebus, Pieris bowdeni, Polyommatus admetus) have not been noted since the end of 19^th century. Moreover, for eleven species: Muschampia baeticus, Agriades optilete, Polyommatus escheri, Tarucus theophrastus, Satyrium lunulata, Melitaea alatauica, M. casta, Hyponephele amardaea, H. comara, Melanargia hylata and M. teneates no information confirming their occurrence in Georgia is available (Table 2) while over twenty other species (recorded mainly by Didmanidze) require confirmation or verification of the identifications.

Table 3:

Comments on new and doubtful butterfly species of Georgia.

#	№	Species name	Comments
1	7.	Spialia orbifer (Hübner, 1823)	Didmanidze (2004) reported this species as Spialia sertorius orbifer (Hübner, 1823). According to the latest revisions of the Spialia genus (Hernández-Roldán et al., 2016), the sertorius species complex includes five species: S. orbifer (Hübner, 1823) distributed in Eastern Europe and Asia, the parapatric S. sertorius (Hoffmannsegg, 1804) occuring in Western Europe and north-western Africa, S. ali (Oberthür, 1881), S. therapne (Rambur, 1832) and S. rosae Hernández-Roldán, Dapporto, Dincă, Vicente & Vila, 2016. We consider that old literature records of S. sertorius for the Caucasus refer to S. orbifer.
2	8.	Favria cribrellum (Eversmann, 1841)	Species reported by Didmanidze (2004), based on only two records. Tshikolovets & Nekrutenko (2012) suggest that this was probably a misidentification, although the species is present in the North Caucasus and may also be found in the northern part of Georgia. Consequently, those records need verification.
3	10*.	Muschampia baeticus (Rambur, 1839)	Species reported only by Didmanidze (2004), not confirmed by other authors and not mentioned by Tshikolovets & Nekrutenko (2012). The closest known localities for this species are in Turkey and Iran (Kudrna et al., 2011). Probably a misidentification.
4	11.	Muschampia poggei (Lederer, 1858)	Species known so far only from areas south of the Lesser Caucasus such as Syria, Armenia, Azerbaijan or Iran (Korb & Bolshakov, 2016). Similar to Tshikolovets & Nekrutenko (2012), we believe that records provided by Didmanidze (1976, 2004) from Georgia require confirmation.
5	20.	Pyrgus carthami (Hübner, 1813)	Recorded by Romanoff (1884), but it is probably a misidentification (Tshikolovets & Nekrutenko, 2012). Didmanidze (2004) reported this species as Pyrgus fritillarius Poda, 1761 considered as a synonym of P. carthami. The closest confirmed localities for this species are in the extreme north western part of the Great Caucasus (Tshikolovets & Nekrutenko, 2012). Consequently, all records from Georgia require verification.
6	21.	Carterocephalus palaemon (Pallas, 1771)	Species known from very old data and only one location: Borjomi (Romanoff, 1884). It has never been recorded again in Georgia. It is impossible to confuse this species with any other, so it is likely to be extinct.
7	36.	Papilio alexanor Esper, 1800	Species not previously reported in Georgia. A single specimen has been found close to David Gareja monastery in K’akheti (Viter, 2022).
8	38. 39.	Leptidea juvernica Williams, 1946 Leptidea sinapis (Linnaeus, 1758)	After the molecular studies on wood white Leptidea butterflies, a new species has been described (Dincă et al., 2011). The oldest known name assigned to the new species is juvernica (Williams, 1946), described as a subspecies of Irish populations of L. reali (Mazel, 2001). In the territory of Georgia, both species may occur: previously described L. sinapis (Linnaeus, 1758) and new species, L. juvernica Williams, 1946. It is impossible to distinguish which old records of L. sinapis refer to L. sinapis or L. juvernica.
9	48.	Pieris mannii (Mayer, 1851)	Species reported only by Didmanidze (2004) and not included by Tshikolovets & Nekrutenko (2012) so it requires confirmation. The closest known localities for this species are in the Republic of Dagestan (Russia, North Caucasus) (Mamedova, 2013).
10	53.	Pieris bowdeni Eitschberger, 1984	Known only from a single specimen from the Zoological Museum in Kyiv; requires confirmation. The closest known localities for this species are in Armenia (Tshikolovets & Nekrutenko, 2012).
11	55.	Pontia edusa (Fabricius, 1777)	P. edusa (Fabricius, 1777) has been separated from its sibling species P. daplidice (Linnaeus, 1758) based on enzyme electrophoretic (Geiger & Scholl, 1982; Geiger, Descimon & Scholl, 1988; Wagener, 1988) and mtDNA studies (John et al., 2013). It is generally known that P daplidice is recorded mainly in Western Europe, North Africa and Cyprus, while P. edusa occurs in the Italian Peninsula, Sicily and Eastern Europe (Dennis et al., 2008; John et al., 2013; Vodă et al., 2016). There were no genetic studies on these species in the Caucasus, but Anikin, Sachkov & Zolotuhin (1993) suggested that P. edusa inhabits the Volga River valley, Crimea, Middle Asia and the Caucasus, while P. daplidice occurs to the north and east of the Rovno-Kiev-Kaluga-N.Novgorod-Chelyabinsk line till Primorye. Based on the distribution maps given in those publications, we assume that all old literature records of P. daplidice from Georgia should be assigned to P. edusa, but this requires corroboration.
12	59.	Colias chrysotheme (Esper, 1781)	A relatively large number of records from Georgia collected by Didmanidze are probably incorrect or doubtful and require confirmation due to the possibility of confusion of this species with other species of the genus Colias. Moreover, Didmanidze (2004) reported this species near Batumi, referring to the publication by Filipjev (1916). In the indicated article, only Colias edusa F. [sic!] is listed in this area, and there are no data about the species C. chrysotheme. C. edusa is a synonym of C. croceus (Geoffroy in A.F. Fourcroy, 1785). The closest confirmed localities for C. chrysotheme are on the northern slopes of the Great Caucasus (Tshikolovets & Nekrutenko, 2012). Due to these ambiguities, all records from Georgia except Abkhazia require verification, because the material is probably misidentified.
13	63.	Colias alfacariensis Ribbe, 1905	Currently, C. alfacariensis and C. hyale (Linnaeus, 1758) are treated as sister species (see Brunton, 1998; Cleary, Descimon & Menken, 2002). It is almost impossible to make a proper identification based on collected adults due to the large variation in wing patterns and other morphological features. C. hyale was reported from the Caucasus region by several different authors, including by Didmanidze (2004), but following the publications by Brunton (1998) and Cleary, Descimon & Menken (2002), all those records should be regarded as C. alfacariensis.
14	72.	Lycaena candens (Herrich-Schäffer, 1844)	Beuret (1952) discovered that Lycaena hippothoe (Linnaeus, 1761) could be divided into two different species: L. hippothoe occurring from East Asia to Spain and L. candens inhabiting a small region from Iran to the Balkans. Based on the distribution maps, all records originally provided by Didmanidze (2004) as L. hippothoe in our work have been assigned to L. candens (Beuret, 1954).
15	74.	Lycaena thetis Klug, 1834	Didmanidze (2004) reported this species in Vashlovani National Park (K’akheti). The closest known localities for this species are in Southern Armenia and Southern Azerbaijan (Snegovaya & Petrov, 2019). Similar to Tshikolovets & Nekrutenko (2012), we believe that those records are misidentified.
16	75.	Lycaena japhetica (Nekrutenko & Effendi, 1983)	For decades, Lycaena (Athamanthia) japhetica was treated as a species from the Caspian area of Eastern Azerbaijan (Tuzov et al., 2000). In 2017, the species was found more than 400 km from known populations, in the vicinity of Kaspi town in Georgia (Morgun, 2019).
17	80*.	Tarucus theophrastus (Fabricius, 1793)	Distributed in the Afrotropical Realm (Deghiche-Diab & Deghiche, 2021). All records for this species reported by Didmanidze (2004) are doubtful and require verification.
18	87.	Pseudophilotes vicrama (Moore, 1865)	Didmanidze (2004) reported two species from the genus Pseudophilotes: P. baton (Bergsträsser, 1779) and P. vicrama. Unfortunately, P. baton is not present in the Caucasus and Transcaucasia. Individuals collected by Didmanidze probably belong to P. vicrama or P. bavius (Eversmann, 1832). The closest known localities for P. bavius are in the Northern Caucasus (Russia) and in Armenia (Tshikolovets & Nekrutenko, 2012), but the species has not been reported in Georgia so far. In agreement with Tshikolovets & Nekrutenko (2012), we believe that the records from Georgia require verification.
19	88.	Scolitanides orion (Pallas, 1771)	Reported by Didmanidze (2004) from only one locality in Georgia (Khelvachauri, close to Batumi) but not confirmed by any other authors. A very distinctive species, impossible to confuse with any other. Species reported from Artvin Province in Turkey, close to Batumi (Karaçetin & Welch, 2011). Record requires confirmation.
20	98.	Plebejus christophi (Staudinger, 1874)	Known only from two localities in Georgia: Aspindza and Khando (Didmanidze, 1979). The closest confirmed localities for this species are in Southern Armenia and Azerbaijan. Records from Georgia require confirmation (Tshikolovets & Nekrutenko, 2012).
21	99. 100. 101.	Kretania modica (Verity, 1935) Kretania sephirus (Frivaldszky, 1835) Kretania stekolnikovi (Stradomsky & Tikhonov, 2015)	After a molecular study on the Kretania pylaon species group, three distinct species have been separated: K. sephirus (Frivaldszky, 1835), K. modica Vérity, 1935 and K. stekolnikovi Stradomsky & Tikhonov, 2015 (Stradomsky & Tikhonov, 2015). All records originally assigned by Didmanidze (2004) to K. sephirus have been divided in our work into those three species based on the distribution maps presented in the above-mentioned publication.
22	110*.	Agriades optilete (Knoch, 1781)	Records provided by Didmanidze (2004) are doubtful and require verification. Based on available literature, this species has not been observed in the Caucasus. The closest known localities for that species are in Eastern Ukraine, the Volga region in Russia and Kazakhstan (Korb & Bolshakov, 2011b, 2016).
23	111.	Agriades dardanus (Freyer, 1843)	Following Higgins & Riley (1970), Didmanidze (2004) reported this species as Agriades glandon dardanus Freyer, 1944. According to Nekrutenko (1974) A. glandon dardanus should be considered as a subspecies of A. pyrenaicus. Unfortunately, this publication was omitted by Didmanidze. Taking the phylogenetic study by Talavera et al. (2013) into account, all recorded specimens should belong to A. pyrenaicus dardanus. Moreover, the aforementioned subspecies was elevated to species level (Wiemers et al., 2018).
24	113.	Lysandra corydonius (Herrich-Schäffer, 1852)	Didmanidze (2004) reported this species as Lysandra corydon (Poda, 1761) following the publication by Jachontov (1914), where it was reported as L. corydon ciscaucasica. The subspecies has been synonymised with Polyommatus corydonius ciscaucasicus by Schurian (1988). P. corydon has not been reported in Georgia (Tshikolovets & Nekrutenko, 2012). The closest known locality for Polyommatus (Lysandra) corydon is on the extreme northern slopes of the Great Caucasus (Lvovsky & Morgun, 2007).
25	115*.	Polyommatus escheri (Hübner, 1823)	The species distribution extends from Southern Europe to Northern Africa. The closest known locality for P. escheri is the European part of Turkey (Karaçetin & Welch, 2011). Two records given by Didmanidze (2004) are doubtful and require verification.
26	119.	Polyommatus icarus (Rottemburg, 1775)	Didmanidze (2004) reported P. icarus and P. persica Bienert, 1870 as two separate species. Curently P. persica is a subspecies of P. icarus (Vodolazhsky & Stradomsky, 2008) and all old records of P. persica have been assigned to P. icarus.
27	122.	Polyommatus admetus (Esper, 1783)	Only two known historical records from Georgia, which were not included by Tshikolovets & Nekrutenko (2012). The closest known localities for this species are in Southern Armenia and Southern Azerbaijan (Tshikolovets & Nekrutenko, 2012). It is highly likely that either that the specimens were misidentified or this species no longer occurs in Georgia.
28	126.	Polyommatus wagneri (Forster, 1956)	Didmanidze (2004) reported this species as Polyommatus damone wagneri (Forster, 1956). Meanwhile, the subspecies was elevated to species level (Lukhtanov et al., 2023).
29	127.	Polyommatus altivagans (Forster, 1956)	A few records from K’akheti in Georgia (Didmanidze, 2004), which were not included by Tshikolovets & Nekrutenko (2012). A relatively large number of records from Armenia, Azerbaijan and Dagestan (Tshikolovets & Nekrutenko, 2012) may suggest that this species could also be found in other regions of Georgia.
30	128.	Polyommatus cyanea (Staudinger, 1899)	The taxonomic position of this species (and subspecies) has changed several times since the studies on the group which were conducted mainly by Forster (1956, 1960, 1961), which described new subspecies within the species Agrodiaetus xerxes. Based on the publication by Forster (1956), Didmanidze (2004) reported this species as Polyommatus xerxes pseudocyanea. Decades later, the species “xerxes” has been considered as subspecies of cyanea within damon-group (Eckweiler & Häuser, 1997).
31	129.	Polyommatus poseidon (Herrich-Schäffer, 1851)	The presence of this species in Georgia has been proven based on a new method using chromosomal and molecular markers (see Lukhtanov & Tikhonov, 2005).
32	132.	Polyommatus aserbeidschana (Forster, 1956)	Initially, it was suspected that this species only occurs in Azerbaijan but, in recent times, it has been confirmed in Iran (Eckweiler & Häuser, 1997), Turkey (Karaçetin & Welch, 2011) and Armenia (Lukhtanov & Dantchenko, 2017). Recently, the species has also been found in Georgia in the Trialetsky Ridge (situated within the Samtskhe-Javakheti region) and the Meskhetsky Ridge (located in the Lesser Caucasus) (Dantchenko, 2000).
33	133.	Polyommatus ninae (Forster, 1956)	The closest known localities for this species are in Southern Armenia and Southern Azerbaijan (Tshikolovets & Nekrutenko, 2012). One record from Tzarskije Kolodtzy (currently known as Dedoplistskaro) in the vicinity of Tbilisi (Forster, 1956) requires verification, but it is probably a misidentification.
34	135.	Afarsia morgiana (Kirby, 1871)	Didmanidze (2004) reported this species as Vacciniina hyrcana Lederer, 1869 following Lewis (1974) nomenclature. Two records from Tbilisi and Shulaveri require confirmation. The closest known localities for this species are in Southern Armenia and Southern Azerbaijan (Snegovaya & Petrov, 2019).
35	138.	Satyrium abdominalis (Gerhard, 1850)	Didmanidze (2004) reported this species as Satyrium acaciae abdominalis (Gerhard, [1850]). The status of this taxon is ambiguous. Some lepidopterologists consider it as a bona species, S. abdominalis, while others regard it as a subspecies of S. acaciae. Similar to Rajaei et al. (2023) in this article, we support the position of this taxon as a valid species.
36	143*.	Satyrium lunulata (Erschoff, 1874)	Single record from Kakheti (Didmanidze, 1979) which is doubtful and requires verification. The closest known localities for this species are in Iran, Tajikistan and Uzbekistan (Tuzov et al., 2000).
37	144.	Callophrys mystaphia Miller, 1913	Single record from Vashlovani National Park (K’akheti) (Didmanidze, 2004) which requires verification. The closest known localities for this species are in Azerbaijan, Iran and Turkey (Krupitsky & Kolesnichenko, 2013).
38	145.	Callophrys chalybeitincta Sovinsky, 1905	Didmanidze (2004) reported this species as Callophrys rubi chalybeitincta Sovinsky, 1905. The subspecies was elevated to species level (Stradomsky & Vodolazhsky, 2011). C. rubi (Linnaeus, 1758) is not present in Georgia.
39	154.	Boloria caucasica (Lederer, 1852)	Didmanidze (2004) reported this species as Boloria pales caucasica Staudinger, 1861. The subspecies was elevated to species level (Simonsen et al., 2010).
40	155.	Boloria eunomia (Esper, 1800)	Single historical record from Akhaltsikhe (Samtskhe-Javakheti) (Bohatsch, 1886) which requires confirmation.
41	158.	Boloria selene (Denis & Schiffermüller, 1775)	Species known from a large number of records listed only by Didmanidze (2004), described as a common species but not confirmed by any other authors. All records and specimens of this species require verification. The closest known localities for Boloria selene are in Armenia (Tshikolovets & Nekrutenko, 2012).
42	169.	Thaleropis ionia (Fischer de Waldheim & Eversmann, 1851)	Species not previously reported from Georgia. The single specimen was found close to Kvariati village in Ajaria (Global Biodiversity Information Facility, 2022).
43	175.	Melitaea interrupta Kolenati, 1846	Didmanidze (2004) reported this species as Melitaea transcaucasica Turati, 1919. In a few other publications based on old data recorded by Didmanidze from the Caucasus, this species is still presented as M. transcaucasica, but currently it is known as a synonym of M. interrupta (Kemal & Koçak, 2013).
44	177*.	Melitaea casta (Kollar, 1848)	Endemic species of Iran (Rajaei et al., 2023). The only one record listed by Didmanidze (2004) is doubtful and requires confirmation.
45	182*.	Melitaea alatauica Staudinger, 1881	Endemic species of the Dzungarian Alatau in south-eastern Kazakhstan (Zhdanko, 2005; Lukhtanov, Gagarina & Pazhenkova, 2021). Records listed by Didmanidze (2004) are not confirmed in any other available resources; probably a misidentification.
46	185.	Euphydryas discordia Bolshakov & Korb, 2013	During the study on females from the nominative subspecies of Euphydryas aurinia (Rottemburg, 1775) from many regions of the Caucasus, a new species to science, E. discordia, was described. It is a cryptic species of E. aurinia group (Bolshakov & Korb, 2013).
47	186.	Euphydryas orientalis (Herrich-Schäffer, 1851)	Only one record from Zekari Pass (Imereti) reported by Didmanidze (2004); it requires verification. The closest known localities for this species are in Central Turkey (Korb & Bolshakov, 2011a).
48	199.	Coenonympha tullia Müller, 1764	Records from Bakuriani and Tskhra-Tskaro reported by Didmanidze (2004) are misidentified (Tshikolovets & Nekrutenko, 2012). The mentioned authors provided a relatively large number of records from Karachay-Cherkessia in Russia. We suspect that this species may occur in Abkhazia.
49	213*.	Hyponephele amardaea (Lederer, 1869)	The closest known localities for this species are in Iran and Turkmenistan (Tuzov et al., 1997). Records listed by Didmanidze (2004) are not confirmed in any other available resources. Probably a misidentification.
50	214*.	Hyponephele comara (Lederer, [1870])	The closest known localities for this species are in Iran and Southern Azerbaijan (Tshikolovets & Nekrutenko, 2012). There is only one record from Khando (Didmanidze, 2004); it requires confirmation. Similar to Tshikolovets & Nekrutenko (2012), we believe this is a misidentification.
51	217*.	Melanargia teneates (Ménétriés, 1832)	Didmanidze (2004) reported this species from Svaneti referring to the publication by Romanoff (1884). In fact, Romanoff gives locality name as “Souant” (arid part of the Talysh Mts., Azerbaijan). It can be assumed that Didmanidze considered that “Souant” means “Svaneti” (in Georgia) or the name was mistyped. Unfortunately, Romanoff made a typographical error in the locality name while referring to a publication by Ménétriés (1832). Ménétriés has provided the correct name for the locality as “Zouvant” (region in the Lesser Caucasus at the Iranian border close to the district town of Lerik). Moreover, Didmanidze provided two own records from Georgia (Svaneti) that have been deemed misidentifications (Tshikolovets & Nekrutenko, 2012).
52	219*.	Melanargia hylata (Ménétriés, 1832)	Single erroneous record from Sukhumi in Abkhazia (Didmanidze, 2004). The author listed this species citing the publication by Romanoff (1884). In fact, Romanoff gives locality name as “Schakouh” in the Talysh Mountains (Azerbaijan/Iran). Didmanidze’s indication of a locality for this species as “Sukhumi” (in Abkhazia) is an error. A locality for this species in the Talysh Mountains was subsequently confirmed by Christoph (1886).

DOI: 10.7717/peerj.18720/table-3

Note:

(#) Comment number used in column [c] in Table 2. (№) Species index used in Table 2.

A complete listing of taxa with known subspecies is available in Table S1 in Supplemental Material.

In total, 7,695 records were found during the study periods, including 3,849 extracted from GBIF. Among them, 7,180 have locality names or geographical coordinates that indicate precise location. In examining the number of records across each period, an upwards trend was apparent. The number of records was the lowest (822) in the first historical period (up to 1945), while in the second period 2,381 records were reported. The highest number of records (4,492) was noted in the last period (post-1991) (Fig. 2). The increased number of records in the last period was mainly due to large scale user activity on Internet platforms like iNaturalist or GBIF.

Figure 2: Number of records collected in every year in three defined periods (see Materials and Methods).

Download full-size image

DOI: 10.7717/peerj.18720/fig-2

List of comments on new and doubtful species

After an examination of all available publications dealing with Georgian butterflies, we conclude that the presence of several dozen species in Georgia is not certain, while several other species reported earlier need confirmation or further systematic review. Since Didmanidze’s checklist (Didmanidze, 2004), several faunistic publications and taxonomic studies incorporating DNA-based techniques have appeared. Hereafter, we have added comments to the species whose status rank has changed due to new genetic research, as well as species whose occurrence in Georgia is doubtful due to the availability of conflicting or ambiguous information/data, misidentifications, or a long-term lack of new records (Table 3).

Distribution, habitat, phenology of Georgian butterflies

Most of recorded species are considered as widely distributed and common in both Georgia and the Caucasus. Twelve species: Muschampia cribrellum, Pyrgus cacaliae, P. carthami, P. serratulae, Euphydryas iduna, Limenitis populi, Coenonympha tullia, Aricia teberdina, Cupido alcetas, Kretania sephirus, K. stekolnikovi and Polyommatus eros are confined only to the Great Caucasus, while sixteen: Colias caucasica, Pieris bowdeni, Melitaea ornata, Coenonympha symphita, Hipparchia fatua, Pseudochazara beroe, P. geyeri, P. mniszechii, Lycaena asabinus, Kretania modica, Polyommatus admetus, P. cyaneus, P. ninae, P. aserbeidschana, P. turcicus, P. damon are restricted to the Lesser Caucasus. Seven species: Eogenes alcides, Muschampia poggei, Papilio alexanor, Afarsia morgiana, Polyommatus altivagans, Tomares romanovi, Hyponephele comara may be found only in the Djavakheti-Armenian plateau. The highest number of butterfly species are found in July, and almost half of the species are strictly univoltine (Table 4).

We created and provided the first list of Georgian vernacular names for butterfly species, which is as complete as possible (see Table 4). Such a list has not yet been published in any article. Creating this list, in our opinion, was very important out of respect for the traditional native names. We extended this list by supplementing it with Russian names, because the Russian language was used in Georgia during the Soviet times, serving as a lingua franca and a language of inter-ethnic communication. Furthermore, since many young people use English, we also added English names. We hope that a list of vernacular names provides useful guides to identify butterflies for the wider entomological community.

The distribution of numerous species is still poorly known because many regions of Georgia have not been explored extensively. Undoubtedly the best studied regions are: Samtskhe-Javakheti (1,713 records, 191 species) and Mtskheta-Mtianeti (1,245 records, 145 species). Some regions, such as K’akheti, Kvemo Kartli and Abkhazia are also well investigated (more than 460 records), with more than 130 species found in them. It is noteworthy that in Samegrelo-Zemo Svaneti and Ajaria, despite having more than 790 records, less than 100 species were noted. The smallest number of records and species were collected in Guria (only 44 records and 30 species) which is really surprising (Table 5, Fig. 3). A relatively large number of records collected from neighbouring regions and direct access to the Black Sea coast suggests that the real number of species in this region is probably much higher than presented.

Table 5:

Comparison of number of records, localities and butterfly species in Georgia regions.

Region	Code¹	Records	Localities	Species
Abkhazia	AB	461	30	131
Ajaria	AJ	794	56	90
Guria	GU	44	12	30
Imereti	IM	316	37	106
K’akheti	KA	924	85	159
Kvemo Kartli	KK	626	70	151
Mtskheta-Mtianeti	MM	1,245	96	145
Rach’a-Lechkhumi-Kvemo Svaneti	RL	105	25	43
Samegrelo-Zemo Svaneti	SZ	808	74	98
Samtskhe-Javakheti	SJ	1,713	81	191
Shida Kartli	SK	100	15	70
Tbilisi	TB	559	11	119
Total in Georgia		7,695	592	244

DOI: 10.7717/peerj.18720/table-5

Note:

1 ISO 3166-2 region code.

Figure 3: Map of studied localities in UTM projection (grid 10 × 10 km).
(˚) One or two record(s). (•) Three or more records. Map credit: Krzysztof Jonko.

Download full-size image

DOI: 10.7717/peerj.18720/fig-3

The arrangement of sampling points on the map (Fig. 3) indicates that the Lesser Caucasus is relatively well explored in comparison to the Great Caucasus. The Great Caucasus, although it borders Lesser Caucasus, remains poorly investigated with regard to butterflies. More species may be expected in this area, but it will depend on the intensity of the future field research.

Discussion

The territory of Georgia is located between two mountain ranges: the Great Caucasus to the north and the Lesser Caucasus to the south. The inter-montane plains of Central Georgia are located between them. It is necessary to remember that a major part of this country is occupied by mountains with 55% of its surface area located at altitudes higher than 1,000 m. Such topographic heterogeneity and habitat diversity favours high species diversity and endemism visible in different taxonomical groups in the entire area of the Caucasus (e.g., Eliava et al., 2007; Sendra & Reboleira, 2012; Zazanashvili et al., 2012; Mumladze, Cameron & Pokryszko, 2014; Konstantinov & Simov, 2018; Grego et al., 2020; Kokhia & Golovatch, 2020).

After including information given by Tshikolovets & Nekrutenko (2012), we updated the list of Georgian butterflies by adding 39 species which increases the total number of species to 244 (belonging to five families). Quite a large number of species new to Georgia compared to the publication mentioned above is a result of three factors: changes in systematics, new discoveries and description of species new to science. It is important to note that, to date, no species have been identified as endemic to Georgia. However, what distinguishes the Georgian butterfly fauna is the presence of rare and extremely rare species (comprising almost 25% of the butterfly fauna), as well as the species endemic to the Caucasus. Our data shows that butterfly fauna is very diverse, despite the fact that Georgia is a relatively small country. It is worth mentioning that Georgia is located at a biogeographical crossroads where at least three biogeographical provinces converge. For this reason, butterfly fauna of this country expresses the mixed character of its biogeographic connections with European, Asian and Middle Eastern elements, as well as Caucasian endemics. Notably, the presence of 16 endemic species to the Caucasus occurring in Georgia indicates a relatively high level of endemism in this country. It is important to note that these species play a crucial role in maintaining biodiversity, not only in Georgia but throughout the entire Caucasus region. Due to their limited geographic range, endemic species are often more vulnerable to extinction. This makes their conservation a priority in light of habitat loss and climate change.

Many records of over a dozen species in Georgia are probably erroneous, although these species can be expected there as they are noted in the surrounding countries. We assumed that the number of species reported in Georgia and its neighbouring countries was comparable.

Consequently, we compared our list of the butterfly fauna with existing checklists (or papers) from neighbouring countries of Armenia and Azerbaijan (belonging to the Transcaucasia) as well as Russia and Turkey (Tshikolovets & Nekrutenko, 2012; Koçak & Kemal, 2018; Langourov, 2019; Sinev, 2019; Snegovaya & Petrov, 2019). In the case of Russia, we focused only the regions bordering Georgia: Krasnodar and Stavropol Krai, the Republic of Adygea, Kabardino-Balkaria, Karachay-Cherkess, North Ossetia-Alania, Ingushetia, referred to as the West Caucasian region and Chechen Republic and the Republic of Dagestan referred to as the East Caucasian region. In the case of Turkey, we concentrated only on two provinces—Artvin and Ardahan.

We found that the number of species recorded for Georgia is comparable to the butterfly fauna of other republics of the Transcaucasia, while it is higher in relation to neighbouring provinces of Turkey and Russia (Table 6).

Table 6:

Comparison of number of butterfly species of Georgia with surrounding countries.

Family	Georgia	Azerbaijan	Armenia	Turkey¹	Russia²
Hesperiidae	29	35	31	24	28
Lycaenidae	83	101	87	82	74
Nymphalidae	96	104	88	83	93
Papilionidae	8	10	5	6	7
Pieridae	28	28	27	24	23
Riodinidae	0	1	0	0	1
Total	244	279	238	219	226

DOI: 10.7717/peerj.18720/table-6

Notes:

1 Only Artvin & Ardahan provinces of Turkey.

2 Only East & West Caucasus regions of Russia.

The high number of species recorded for Azerbaijan should be treated with caution; we are convinced that several species require independent confirmation, like in the case of Georgia.

Even though Georgia has a long history of butterfly research, the fauna is still poorly researched. It is worth mentioning that, since the collapse of the Soviet Union, research on Georgian butterflies and moths has primarily been conducted by Didmanidze (2004), and it has been 20 years since the last inventory was published.

The updated checklist presented in this article and data containing thumbnails, vernacular names and species regional distribution maps may serve as a baseline for further exploration of Georgian butterflies. We hope that our work will be useful for scientists and will also promote interest in butterflies among communities, local entomologists and citizen scientists. They may use this checklist to document the presence or absence of butterfly species in specific regions of Georgia. Citizen participation in data collection may be one of the steps towards conservation of butterflies and/or their habitats in that country.

Conclusions

Our study supplements the knowledge of butterflies from Georgia. We added 39 new species found or described in the last decade. Among 244 recorded species almost 25% of Georgian species are categorised as rare and extremely rare, and these species may be at the highest risk of potential extinction. The number of recorded species in Georgia is comparable to the butterfly fauna of other republics of the Transcaucasia; however, it is higher than neighbouring provinces of Turkey and Russia. The occurrence of over a dozen species is questioned by us due to potential misidentification or ambiguous data. That is why we consider it essential to verify determination of several species listed by Didmanidze or confirm their occurrence in Georgia with independent studies. The data presented in this article may serve as a baseline for further exploration of the diversity of butterfly fauna of Georgia. This is crucial to better management of conservation in the face of loss habitats and changes resulting from climate change.

Supplemental Information

Complete list of Georgian butterflies with known subspecies.

DOI: 10.7717/peerj.18720/supp-1

Download

Raw data with location names and GPS coordinates used in the study.

DOI: 10.7717/peerj.18720/supp-2

Download

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Abbr	Region name–English	Region name–Georgian	Administrative divisions of Georgia
AB	Abkhazia	აფხაზეთი
AJ	Adjara	აჭარა
GU	Guria	გურია
IM	Imereti	იმერეთი
KA	K’akheti	კახეთი
KK	Kvemo Kartli	ქვემო ქართლი
MM	Mtskheta-Mtianeti	მცხეთა-მთიანეთი
RL	Rach’a-Lechkhumi-Kvemo Svaneti	რაჭა-ლეჩხუმი და ქვემო სვანეთი
SJ	Samtskhe-Javakheti	სამცხე-ჯავახეთი	Table symbols legend:
SK	Shida Kartli	შიდა ქართლი	˚ Historical records (collected up to the 1945)
SZ	Samegrelo-Zemo Svaneti	სამეგრელო-ზემო სვანეთი	• Old records (collected between 1946 and 1991)
TB	Tbilisi	თბილისი	▀ Recent records (collected after 1991)

№	Species name	AB	AJ	GU	IM	KA	KK	MM	RL	SJ	SK	SZ	TB	[c]
	Hesperiidae Latreille, 1802
	Pyrginae Burmeister, 1878
1	Erynnis tages (Linnaeus, 1758)	•	▀	•	▀	▀	▀	▀	•	▀	•	▀	•
2	Erynnis marloyi (Boisduval, 1834)	•				▀	▀						▀
3	Carcharodus alceae (Esper, 1780)	•	▀		•	▀	▀			▀			▀
4	Carcharodus lavatherae (Esper, 1783)						▀	•		˚			˚
5	Carcharodus floccifera (Zeller, 1847)	•				▀	•			•			˚
6	Carcharodus orientalis Reverdin, 1913	•			•		▀				•
7	Spialia orbifer (Hübner, 1823)	•			˚	▀	▀	▀		•			▀	1
8	Favria cribrellum (Eversmann, 1841)						•						˚	2
9	Muschampia tessellum (Hübner, 1803)		•										˚
10*	Muschampia baeticus (Rambur, 1839)	•	•		•					•				3
11	Muschampia poggei (Lederer, 1858)						•	•						4
12	Pyrgus melotis (Duponchel, 1834)	•	˚	•		▀	▀	▀	▀	▀		•	•
13	Pyrgus serratulae (Rambur, 1839)	•			˚	˚	•	▀		•
14	Pyrgus armoricanus (Oberthür, 1910)	▀				▀	•	▀		•		▀	▀
15	Pyrgus alveus Hübner, 1803	•				•	▀	▀		▀		•	˚
16^✠	Pyrgus jupei (Alberti, 1967)		•		▀			▀		▀
17	Pyrgus sidae (Esper, 1784)					▀	▀	•		▀			•
18	Pyrgus cacaliae (Rambur, 1839)	•
19	Pyrgus cinarae (Rambur, 1839)						•	•
20	Pyrgus carthami (Hübner, 1813)					•	˚			•				5
	Heteropterinae Aurivillius, 1925
21	Carterocephalus palaemon (Pallas, 1771)									˚				6
22	Heteropterus morpheus (Pallas, 1771)	•	•		•							•
	Hesperiinae Latreille, 1809
23	Eogenes alcides (Herrich-Schäffer, 1852)					▀							•
24	Gegenes nostrodamus (Fabricius, 1793)					•		▀					•
25	Thymelicus hyrax (Lederer, 1861)						•
26	Thymelicus sylvestris (Poda, 1761)	•			▀	▀		▀		▀	•		•
27	Thymelicus lineola (Ochsenheimer, 1808)	•	▀		▀		▀	▀		▀			▀
28	Ochlodes sylvanus (Esper, 1777)	•	•			▀	▀	▀		▀	•	▀	▀
29	Hesperia comma (Linnaeus, 1758)	▀	▀		▀	▀		▀		˚	▀	▀
	Papilionidae Latreille, 1802
	Parnassiinae Duponchel, 1835
30	Parnassius mnemosyne (Linnaeus, 1758)	•	▀		▀	˚	•			▀		▀	˚
31^✠	Parnassius nordmanni Ménétriés, 1850	•	•	•			•	▀	▀			▀
32	Parnassius phoebus (Fabricius, 1793)		˚
33	Parnassius apollo (Linnaeus, 1758)	•	▀		•	▀		▀	▀	▀		▀	˚
34	Zerynthia caucasica (Lederer, 1864)	•	▀		▀	•			•	˚		•
	Papilioninae Latreille, 1802
35	Iphiclides podalirius (Linnaeus, 1758)	▀	▀		▀	▀	▀	▀		▀	•	▀	▀
36	Papilio alexanor Esper, 1800					▀								7
37	Papilio machaon Linnaeus, 1758	•	▀		▀	▀	▀	▀		▀		▀	▀
	Pieridae Swainson, 1820
	Dismorphiinae Schatz, 1887
38	Leptidea sinapis (Linnaeus, 1758)	•	•		˚	▀	▀	•		▀	•	˚	▀	8
39	Leptidea juvernica Williams, 1946	?	?		▀	▀	?	▀		?	?	?	?	8
40	Leptidea duponcheli (Staudinger, 1871)					▀	▀			▀			▀
	Pierinae Duponchel, 1835
41	Anthocharis cardamines (Linnaeus, 1758)	•	▀			▀	▀	▀		▀	•	•	▀
42	Anthocharis damone Boisduval, 1836						•			▀
43	Anthocharis gruneri Herrich-Schäffer, 1851									˚			▀
44	Zegris eupheme (Esper, 1804)					•	•			•			•
45	Euchloe ausonia (Hübner, 1804)		•		•	▀	▀	▀		▀			▀
46	Aporia crataegi (Linnaeus, 1758)	▀	•			▀	▀	▀	▀	▀	•	▀	▀
47	Pieris brassicae (Linnaeus, 1758)	•	▀	•	▀	▀	▀	▀	•	▀	•	˚	▀
48	Pieris mannii (Mayer, 1851)					•		•	˚					9
49	Pieris rapae (Linnaeus, 1758)	▀	▀		˚	▀	▀	▀		▀		˚	▀
50	Pieris ergane (Geyer, 1828)		•	•		▀				•
51	Pieris napi (Linnaeus, 1758)	•	▀		▀	▀	▀	▀		▀	•		•
52	Pieris bryoniae (Hübner, 1806)	•						▀	▀	▀		▀
53^✠	Pieris bowdeni Eitschberger, 1984									˚				10
54	Pontia callidice (Hübner, 1800)					▀	˚	˚	•	•		•
55	Pontia edusa (Fabricius, 1777)	•	▀	•	•	▀	▀	▀	•	▀	▀	•	▀	11
56	Pontia chloridice (Hübner, 1813)									▀
	Coliadinae Swainson, 1827
57	Colias erate (Esper, 1805)	•	▀		•		•	•		˚	•	•	▀
58	Colias croceus (Geoffroy, 1785)	•	▀	•	▀	▀	▀	▀	•	▀	▀	▀	▀
59	Colias chrysotheme (Esper, 1781)	•	˚		•	•	•		•	•		•		12
60	Colias aurorina Herrich-Schäffer, 1850	˚			˚	▀	•			•			˚
61	Colias caucasica Staudinger, 1871				˚	•	•			▀
62	Colias thisoa Ménétriés, 1832				•	•				•
63	Colias alfacariensis Ribbe, 1905	•	•		•		▀	▀		▀	▀	▀	▀	13
64	Gonepteryx rhamni (Linnaeus, 1758)	•	▀	•	▀	▀	▀	▀	▀	▀	•	▀	▀
65	Gonepteryx farinosa (Zeller, 1847)		•				▀	▀		•			▀
	Lycaenidae Leach, 1815
	Lycaeninae Leach, 1815
66	Lycaena phlaeas (Linnaeus, 1760)	•	▀		▀	▀	▀	▀		▀	•	▀	▀
67	Lycaena helle Denis & Schiffermüller, 1775	•								˚		▀
68	Lycaena dispar (Haworth, 1802)	•		•	▀	▀	▀	•		•	•	▀
69	Lycaena virgaureae (Linnaeus, 1758)	•	•	•	•	▀	▀	▀	▀	▀	•	▀
70	Lycaena tityrus Poda, 1761	▀	•		•	▀	▀	▀	▀	▀		▀	▀
71	Lycaena alciphron Rottemburg, 1775					•	▀	▀		▀			▀
72	Lycaena candens (Herrich-Schäffer, 1844)	▀			˚	▀	•	▀		▀		•		14
73	Lycaena thersamon (Esper, 1784)	•			˚	▀	▀	•		▀		•	▀
74	Lycaena thetis Klug, 1834					•								15
75^✠	Lycaena japhetica (Nekrutenko & Effendi, 1983)										▀			16
76	Lycaena asabinus (Herrich-Schäffer, 1851)					•	•			•
77	Lycaena ochimus (Herrich-Schäffer, 1851)	•					•			•
	Polyommatinae Swainson, 1827
78	Lampides boeticus (Linnaeus, 1767)	•	▀		•	•				▀		▀
79	Leptotes pirithous (Linnaeus, 1767)	▀	•	•	•							•
80*	Tarucus theophrastus (Fabricius, 1793)					•	˚							17
81	Tarucus balkanica (Freyer, 1844)	•				▀	▀	▀					▀
82	Cupido minimus (Fuessly, 1775)	•	▀			•	•	▀		▀		•	•
83	Cupido osiris (Meigen, 1829)				•	•	•	•		•	•		˚
84	Cupido argiades (Pallas, 1771)	•	▀		▀	▀	▀	▀		▀	•	▀	•
85	Cupido alcetas (Hoffmannsegg, 1804)	•
86	Celastrina argiolus (Linnaeus, 1758)	•	▀		▀	▀	▀	▀		▀		▀	▀
87	Pseudophilotes vicrama (Moore, 1865)		•	•		▀	•	▀		▀				18
88	Scolitantides orion (Pallas, 1771)		•											19
89	Glaucopsyche alexis (Poda, 1761)	•	•		•	▀	▀	•		▀			▀
90	Phengaris arion (Linnaeus, 1758)	▀			˚	▀	▀	•		▀	˚
91	Phengaris teleius (Bergsträsser, 1779)									•		•
92	Phengaris nausithous (Bergsträsser, 1779)					•	▀			▀
93	Phengaris alcon (Denis & Schiffermüller, 1775)				•		▀			▀	•
94	Luthrodes galba (Lederer, 1855)					•
95	Plebejus argus (Linnaeus, 1758)	•	▀	•	•		▀	▀	▀	▀	▀	•	•
96	Plebejus idas (Linnaeus, 1761)	•						▀		▀
97	Plebejus argyrognomon (Bergsträsser, 1779)	•	▀		▀			▀		▀	•	▀	•
98	Plebejus christophi (Staudinger, 1874)									•				20
99	Kretania sephirus (Frivaldszky, 1835)	•						▀						21
100	Kretania modica (Vérity, 1935)									▀				21
101	Kretania stekolnikovi Stradomsky & Tikhonov, 2015					▀								21
102	Kretania eurypilus (Freyer, 1851)					•				•
103	Plebejidea loewii (Zeller, 1847)									▀
104	Aricia agestis (Denis & Schiffermüller, 1775)	•			•	▀	▀	▀		▀	▀	▀	▀
105	Aricia artaxerxes (Fabricius, 1793)	•					˚	▀		▀		▀
106	Aricia anteros (Freyer, 1838)				•		▀	▀		▀	•	▀
107	Eumedonia eumedon (Esper, 1780)	▀			˚	▀	•	▀	▀	▀		▀
108	Cyaniris semiargus (Rottemburg, 1775)	•	•			▀	▀	▀		▀		▀	▀
109*	Agriades optilete (Knoch, 1781)						•			•		•		22
110	Agriades dardanus (Freyer, 1843)	•			•	▀		▀		▀		▀		23
111	Lysandra bellargus (Rottemburg, 1775)	•	•			▀	▀	▀		▀	▀	˚	•
112	Lysandra corydonius (Herrich-Schäffer, 1852)	•			•	▀	▀	▀	▀	˚	•	▀		24
113	Neolysandra coelestina (Eversmann, 1843)					▀		•		▀
114*	Polyommatus escheri (Hübner, 1823)	•								•				25
115	Polyommatus dorylas (Denis & Schiffermüller, 1775)				˚		˚	▀		▀
116	Polyommatus amandus (Schneider, 1792)	•			•	▀	▀	▀		▀		▀	˚
117	Polyommatus thersites (Cantener, 1835)		•			▀	▀	▀		▀	•		▀
118	Polyommatus icarus (Rottemburg, 1775)	•	▀	•	▀	▀	▀	▀	•	▀	•	▀	▀	26
119	Polyommatus eros (Ochsenheimer, 1808)	•				•		▀				▀
120	Polyommatus daphnis (Denis & Schiffermüller, 1775)	•	•		˚	•	▀	▀		▀	•	▀	▀
121	Polyommatus admetus (Esper, 1783)									˚			˚	27
122	Polyommatus ripartii (Freyer, 1830)						▀	▀			•
123^✠	Polyommatus eriwanensis (Forster, 1960)	•											•
124	Polyommatus damon (Denis & Schiffermüller, 1775)						•			▀
125^✠	Polyommatus wagneri (Forster, 1956)									•				29
126	Polyommatus altivagans (Forster, 1956)					•								29
127	Polyommatus cyanea (Staudinger, 1899)					•		•		▀				30
128	Polyommatus poseidon (Herrich-Schäffer, 1851)									▀				31
129^✠	Polyommatus teberdina (Sheljuzhko, 1934)	•				•		•				•
130	Polyommatus turcicus (Koçak, 1977)									▀
131	Polyommatus aserbeidschana (Forster, 1956)									▀				32
132	Polyommatus ninae (Forster, 1956)												˚	33
133	Polyommatus demavendi (Pfeiffer, 1938)									•
134	Afarsia morgiana (Kirby, 1871)						•						•	34
	Theclinae Swainson, 1831
135	Satyrium w-album (Knoch, 1782)	•	•			▀			˚	•		•
136	Satyrium spini (Denis & Schiffermüller, 1775)					▀	•			▀			•
137	Satyrium abdominalis (Gerhard, 1850)					▀		•		•				35
138	Satyrium ilicis (Esper, 1779)	•			•	•	•	▀		▀	•		•
139	Satyrium acaciae (Fabricius, 1787)					•	•			•			˚
140	Satyrium ledereri (Boisduval, 1848)					▀	•			▀			▀
141	Satyrium hyrcanica (Riley, 1939)					•
142*	Satyrium lunulata (Erschoff, 1874)					•								36
143^✠	Callophrys mystaphia Miller, 1913					•								37
144	Callophrys chalybeitincta Sovinsky, 1905	•	▀			▀	•	•		•	•	▀	▀	38
145	Thecla betulae (Linnaeus, 1758)	•			•	•				▀
146	Favonius quercus (Linnaeus, 1758)	•			▀	•	▀	•		▀			▀
147	Tomares callimachus (Eversmann, 1848)					▀	•			•			•
148^✠	Tomares romanovi (Christoph, 1882)					▀
	Nymphalidae Rafinesque, 1815
	Limenitidinae Behr, 1864
149	Limenitis populi (Linnaeus, 1758)	•
150	Limenitis camilla (Linnaeus, 1764)	•	•		•		▀	▀		▀	•		▀
151	Limenitis reducta Staudinger, 1901	•	•		˚	•	▀	▀		▀	•	˚	▀
152	Neptis rivularis (Scopoli, 1763)	•				▀	▀	▀		▀			▀
	Heliconiinae Swainson, 1822
153^✠	Boloria caucasica (Lederer, 1852)	•		˚	•			▀	▀	▀		▀		39
154	Boloria eunomia (Esper, 1800)									˚				40
155	Boloria euphrosyne (Linnaeus, 1758)	•	▀	•	•		•	▀		▀		•
156	Boloria selene (Denis & Schiffermüller, 1775)		•	•			•	•						41
157	Boloria dia (Linnaeus, 1767)	•	•	•	•	▀	▀	▀		▀	•	•	˚
158	Issoria lathonia (Linnaeus, 1758)	•	▀	•	•	▀	▀	▀		▀		▀	▀
159	Argynnis paphia (Linnaeus, 1758)	•	▀	▀	▀	▀	▀	▀	▀	▀	▀	▀	▀
160	Argynnis pandora (Denis & Schiffermüller, 1775)	•			•	▀	▀	▀		▀	▀	•	▀
161	Speyeria aglaja (Linnaeus, 1758)	•	•		▀	▀	▀	▀	˚	▀		▀
162	Fabriciana adippe (Denis & Schiffermüller, 1775)	•	•			▀	•	▀	˚	▀		▀	•
163	Fabriciana niobe (Linnaeus, 1758)	•				▀	▀	▀		▀	•		•
164	Brenthis ino (Rottemburg, 1775)					▀	˚			▀
165	Brenthis daphne (Bergsträsser, 1780)	•				▀				▀
166	Brenthis hecate (Denis & Schiffermüller, 1775)					▀	▀	•		▀		▀	▀
	Apaturinae Boisduval, 1840
167	Apatura ilia (Denis & Schiffermüller, 1775)	•			▀		•	▀	▀	•			•
168	Thaleropis ionia Eversmann, 1851		▀											42
	Nymphalinae Swainson, 1827
169	Melitaea cinxia (Linnaeus, 1758)	•	▀		▀	▀	▀	▀		▀		▀	•
170	Melitaea arduinna (Esper, 1783)					•				▀	•		•
171	Melitaea diamina (Lang, 1789)	•			•			▀		▀		▀
172	Melitaea phoebe (Denis & Schiffermüller, 1775)	•	•		˚	▀	▀			•	•		•
173	Melitaea ornata Christoph, 1893						•			•
174	Melitaea interrupta Kolenati, 1846	•			•	▀	▀	▀	▀	▀		▀	•	43
175	Melitaea persea Kollar, 1850					▀	•
176*	Melitaea casta (Kollar, 1848)							•						44
177	Melitaea trivia (Denis & Schiffermüller, 1775)					•		▀		˚
178	Melitaea athalia (Rottemburg, 1775)	•	•		•	▀	▀	•	•	▀			•
179^✠	Melitaea caucasogenita Vérity, 1930							▀		▀	▀	▀	▀
180	Melitaea aurelia Nickerl, 1850									▀
181*	Melitaea alatauica Staudinger, 1881	•								•				45
182	Euphydryas iduna (Dalman, 1816)					•		•
183	Euphydryas aurinia (Rottemburg, 1775)		•		•	▀	▀	▀	▀	▀		▀	•
184	Euphydryas discordia Bolshakov & Korb, 2013									▀				46
185	Euphydryas orientalis (Herrich-Schäffer, 1851)				•									47
186	Vanessa atalanta (Linnaeus, 1758)	▀	▀	▀	▀	▀	▀	▀	▀	▀	•	▀	•
187	Vanessa cardui (Linnaeus, 1758)	•	▀	•	▀	▀	▀	▀	•	▀	•	▀	▀
188	Araschnia levana (Linnaeus, 1758)	•					•
189	Aglais urticae (Linnaeus, 1758)	•	•		▀	▀	▀	▀		▀	•	▀	▀
190	Aglais io (Linnaeus, 1758)	▀	▀	•	•	▀	▀	▀		▀	•	▀	▀
191	Nymphalis antiopa (Linnaeus, 1758)	▀			▀	▀		▀	▀	▀	•	▀	▀
192	Nymphalis polychloros (Linnaeus, 1758)	•	•			▀	•	▀			•	•	▀
193	Nymphalis xanthomelas (Denis & Schiffermüller, 1775)								•	•
194	Nymphalis vaualbum (Denis & Schiffermüller, 1775)	•								˚		▀
195	Polygonia egea (Cramer, 1775)					˚	▀	▀		▀			▀
196	Polygonia c-album (Linnaeus, 1758)	•	▀	•	•	▀	▀	▀	▀	▀	•	▀
	Satyrinae Boisduval, 1833
197	Coenonympha pamphilus (Linnaeus, 1758)	•	•	•	▀	▀	▀	▀	▀	▀	•	•	▀
198	Coenonympha tullia (Müller, 1764)	•												48
199	Coenonympha glycerion (Borkhausen, 1788)					•	•	▀		▀
200	Coenonympha arcania (Linnaeus, 1760)	•	•		▀	•	▀	▀		▀	•		˚
201	Coenonympha leander (Esper, 1784)					•		▀		•		▀
202^✠	Coenonympha saadi Kollar, 1849					•				•
203^✠	Coenonympha symphita Lederer, 1870		•		•	•	•			▀
204	Kirinia climene (Esper, 1783)	•			˚		▀	▀		▀			▀
205	Pararge aegeria (Linnaeus, 1758)	▀	▀		˚	▀	▀	▀	•	▀		•	▀
206	Lasiommata megera (Linnaeus, 1767)	•	▀		▀	▀	▀	▀		▀	•	•	▀
207	Lasiommata petropolitana (Fabricius, 1787)				•	▀	˚	•		▀
208	Lasiommata maera (Linnaeus, 1758)	•	▀		˚	▀	▀	▀	•	▀	•	▀	▀
209	Maniola jurtina (Linnaeus, 1758)	•	▀	•	▀	▀	▀	▀	▀	▀	•	▀	▀
210	Hyponephele lycaon (Kühn, 1774)	•	▀		•	▀	•	▀		▀	•		˚
211	Hyponephele lupinus (Costa, 1836)					▀	▀	•					▀
212*	Hyponephele amardaea (Lederer, 1869)	•						•		•	•			49
213*	Hyponephele comara (Lederer, [1870])									▀				50
214	Melanargia russiae (Esper, 1783)		▀			▀	•	▀	•	▀	•		▀
215	Melanargia galathea (Linnaeus, 1758)	•			▀	▀	▀	▀	•	▀	▀	•	▀
216*	Melanargia teneates (Ménétriés, 1832)						•	•				˚		51
217	Melanargia larissa (Geyer, 1828)				▀	•	▀	▀		▀	•		▀
218*	Melanargia hylata (Ménétriés, 1832)	˚												52
219	Brintesia circe (Fabricius, 1775)			•	▀	•	▀	▀		▀			▀
220	Arethusana arethusa (Denis & Schiffermüller, 1775)					•	˚	˚		▀	•		•
221	Chazara briseis (Linnaeus, 1764)	•			˚	▀	▀	▀	▀	▀	•	•	▀
222	Chazara persephone (Hübner, 1805)					•	˚	˚		•			˚
223	Pseudochazara alpina (Staudinger, 1878)			•		▀	•	˚		•
224	Pseudochazara mniszechii (Herrich-Schäffer, 1851)							•		•		•	•
225	Pseudochazara geyeri (Herrich-Schaffer, 1846)						˚			•
226	Pseudochazara beroe (Herrich-Schaffer, 1844)									•
227	Pseudochazara pelopea (Klug, 1832)	•		˚		•	▀	▀		•			•
228	Satyrus iranica Schwingenschuss, 1939		•		˚	˚	•	•		▀	˚		˚
229	Satyrus amasina Staudinger, 1861					•				•
230	Minois dryas (Scopoli, 1763)	•	▀		▀	•	▀	▀		▀		▀	▀
231	Hipparchia statilinus (Hufnagel, 1766)				˚	▀	•	˚		▀			•
232	Hipparchia fatua Freyer, 1843				•					▀
233	Hipparchia pellucida (Stauder, 1924)	•				▀	▀	▀		▀		˚	▀
234	Hipparchia syriaca (Staudinger, 1871)	•	•		•	▀	•	▀		▀	•	▀	▀
235	Hipparchia autonoe (Esper, 1783)	•
236	Hipparchia parisatis (Kollar, 1849)					▀
237	Erebia aethiops (Esper, 1777)	•	▀		▀	▀	▀	▀	▀	▀	•	▀	˚
238	Erebia medusa (Denis & Schiffermüller, 1775)	•	•			▀	˚	▀		▀	˚	•
239	Erebia iranica Grum-Grshimailo, 1895	•	•		•	•	•	•		▀		•	˚
240^✠	Erebia graucasica Jachontov, 1909	•	•		▀	•	•	•		▀	˚	•
241^✠	Erebia melancholica Herrich-Schäffer, 1846	▀	▀	•	˚	•	▀	▀	▀	▀		▀
242^✠	Erebia hewitsonii Lederer, 1864		▀		▀					▀
243	Proterebia afra (Fabricius, 1787)					▀	•	•			•
	Libytheinae Boisduval, 1833
244	Libythea celtis (Laicharting, 1782)	•				▀				˚

Hesperiidae, Latreille, 1809		Pyrginae, Burmeister, 1878
1. *Erynnis tages* (Linnaeus, 1758)	English name:	Dingy Skipper
	Georgian name:	მოშავო თავმსხვილა
	Russian name:	Бурокрылка Тагес
	Distr. Caucasus:	Widespread.
	Distr. Georgia:	Widespread and common.
	Flight:	Bivoltine in the lowlands, April to August. Univoltine in the mountains, late May to July.
	Habitat:	Various open, grassy habitats.
2. *Erynnis marloyi* (Boisduval, 1834)	English name:	Inky Skipper
	Georgian name:
	Russian name:	Бурокрылка Марлоя
	Distr. Caucasus:	Central and eastern part of the Great Caucasus, Lesser Caucasus.
	Distr. Georgia:	Local and rare.
	Flight:	Uni- or bivoltine, April to October depending on the altitude.
	Habitat:	Dry, stony and clayey places, rocky gorges and slopes.
3. *Carcharodus alceae* (Esper, 1780)	English name:	Mallow Skipper
	Georgian name:	ბალბის თავმსხვილა
	Russian name:	Зубчатокрылка альцея
	Distr. Caucasus:	Widespread.
	Distr. Georgia:	Local and rare.
	Flight:	Multivoltine in the lowlands, April to October. Univoltine in the mountains.
	Habitat:	Meadows, steppes, grassy slopes, woodland clearings.
4. *Carcharodus lavatherae* (Esper, 1783)	English name:	Marbled Skipper
	Georgian name:
	Russian name:	Толстоголовка чистецовая
	Distr. Caucasus:	Widespread.
	Distr. Georgia:	Local, extremely rare.
	Flight:	Univoltine, June to August.
	Habitat:	Dry grassland with rocky calcareous places, dry slopes.
5. *Carcharodus floccifera* (Zeller, 1847)	English name:	Tufted Marbled Skipper
	Georgian name:
	Russian name:	Зубчатокрылка шандровая
	Distr. Caucasus:	Widespread.
	Distr. Georgia:	Widespread but not frequent.
	Flight:	Bivoltine in the lowlands, May to August. Univoltine in the high mountains, June to August.
	Habitat:	Meadows, grassy slopes, woodland clearings.
6. *Carcharodus orientalis* Reverdin, 1913	English name:	Oriental Marbled Skipper
	Georgian name:
	Russian name:	Толстоголовка восточная
	Distr. Caucasus:	Widespread, except Colchis and Kura lowlands.