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An updated checklist of the European Butterflies (Lepidoptera, Papilionoidea)
expand article infoMartin Wiemers, Emilio Balletto§, Vlad Dincă|, Zdenek Faltynek Fric, Gerardo Lamas#, Vladimir Lukhtanov¤, Miguel L. Munguira«, Chris A. M. van Swaay», Roger Vila˄, Albert Vliegenthart», Niklas Wahlberg˅, Rudi Verovnik¦
‡ UFZ – Helmholtz Centre for Environmental Research, Halle, Germany
§ Turin University, Torino, Italy
| University of Oulu, Oulu, Finland
¶ Biology Centre CAS, Ceske Budejovice, Czech Republic
# Universidad Nacional Mayor de San Marcos, Lima, Peru
¤ Zoological Institute of Russian Academy of Sciences, St. Petersburg, Russia
« Universidad Autónoma de Madrid, Madrid, Spain
» Dutch Butterfly Conservation, Wageningen, Netherlands
˄ CSIC-Universitat Pompeu Fabra, Barcelona, Spain
˅ Lund University, Lund, Sweden
¦ University of Ljubljana, Ljubljana, Slovenia
Open Access

Abstract

This paper presents an updated checklist of the butterflies of Europe, together with their original name combinations, and their occurrence status in each European country. According to this checklist, 496 species of the superfamily Papilionoidea occur in Europe. Changes in comparison with the last version (2.6.2) of Fauna Europaea are discussed. Compared to that version, 16 species are new additions, either due to cryptic species most of which have been discovered by molecular methods (13 cases) or due to discoveries of Asian species on the eastern border of the European territory in the Ural mountains (three cases). On the other hand, nine species had to be removed from the list, because they either do not occur in Europe or lost their species status due to new evidence. In addition, three species names had to be changed and 30 species changed their combination due to new evidence on phylogenetic relationships. Furthermore, minor corrections were applied to some authors’ names and years of publication. Finally, the name Polyommatus ottomanus Lefèbvre, 1831, which is threatened by its senior synonym Lycaena legeri Freyer, 1830, is declared a nomen protectum, thereby conserving its name in the current combination Lycaena ottomana.

Keywords

checklist, butterflies, Europe

Introduction

Butterflies constitute one of the best-known groups of insects and have become important models to study speciation, community ecology, biogeography, climate change, and insect-plant interactions. With close to 19,000 described species [18,768 presumably valid species recorded by 2011; that figure is higher today, i.e., ca. 19,000 species], they represent about 12% of currently known species of Lepidoptera (Van Nieukerken et al. 2011). According to current molecular systematics (Mutanen et al. 2010; Heikkilä et al. 2012; Espeland et al. 2018), the single butterfly superfamily Papilionoidea comprises 7 families (Table 1, Fig. 1) and includes the Hesperiidae (skippers) and Hedylidae (moth butterflies). The skippers have previously been thought to represent the sister group to the butterflies and were often placed in a separate superfamily Hesperioidea, but the molecular results indicate that the family Papilionidae is the sister to the remaining butterflies, which also include the small Neotropical family Hedylidae with only 36 species. Apart from the latter family, all butterfly families are represented on all continents except Antarctica, although most species of Riodinidae are confined to the Neotropical Region. Butterfly diversity is particularly high in the tropics, especially the Neotropics, and only 496 species are found in Europe according to the present checklist.

Family systematics of butterflies.

Superfamily Papilionoidea Latreille, [1802] Genera* Species*
Family Papilionidae Latreille, [1802] 32 570
Family Hedylidae Guenée, [1858] 1 36
Family Hesperiidae Latreille, 1809 570 4113
Family Pieridae Swainson, 1820 91 1164
Family Riodinidae Grote, 1895 146 1532
Family Lycaenidae [Leach], [1815] 416 5201
Family Nymphalidae Rafinesque, 1815 559 6152
Figure 1. 

Global species richness of butterfly families.

The taxonomy of butterflies started in 1758 with the Swedish naturalist Carl von Linné (Latinised to Carolus Linnaeus), who introduced binominal nomenclature and described the highest number of European butterfly species, all of them in a single genus Papilio. Seventy-one of them currently still hold the names given by Linné, albeit mostly in different genera. Other authors who described many new species during the 18th century were the German entomologists Eugen Johann Christoph Esper and Jacob Hübner, the Danish entomologist Johann Christian Fabricius, as well as the Austrian lepidopterist Johann Ignaz Schiffermüller (the latter in an anonymous publication usually referred to as [Denis & Schiffermüller], but see Kudrna and Belicek (2005), Sattler and Tremewan (2009) and Kudrna (2015) for a controversial debate on this topic). By 1820, half of the European butterfly fauna had been validly described, and species were placed in a growing number of genera (starting with Hesperia Fabricius, 1793 as the second-named genus for the skippers). During the 19th century, more than 60 European lepidopterists continued the inventory of Europe’s butterfly fauna, and the first overview of Palearctic butterflies (and other Lepidoptera) was published by Seitz (1907–1909). At that time, already 90% of Europe’s butterfly species had been described and the rate of newly discovered species slowed down (Fig. 2). Another milestone for butterfly research in Europe was the field guide of Higgins and Riley (1970), which included distribution maps of Western Palearctic butterflies, and led to a growing interest in butterflies across Europe. This field guide was also translated into other languages (e.g., German, French, and Spanish) and updated several times (most recently by Tolman and Lewington 2008). However, despite their somehow misleading titles, these guides excluded large parts of eastern Europe (i.e., Belarus, Ukraine, Moldova and most of Russia (apart from Kaliningrad enclave) and therefore all the species from the Ural mountains). The proliferation of butterfly field guides by various authors across Europe also led to an increasing confusion of butterfly nomenclature due to different taxonomic concepts. The first step to standardize European butterfly taxonomy and the precursor of our list was the book (and accompanying CD) by Karsholt and Razowski (1996). It constituted a country-level checklist of all European Lepidoptera, but excluding the Mid-Atlantic islands (i.e., Canary Islands, Madeira, and Azores) and contained 440 butterfly species. This book was also the basis for the list of Lepidoptera in the online database Fauna Europaea, a project under the auspices of the European Commission, which started in 2000 (De Jong et al. 2014) and aimed to provide checklists for all European animal species. This database, which went online on 16 December 2004, also included Cyprus and the Mid-Atlantic islands, which are hotspots of narrow endemics. At about the same time, the first distribution atlas of all European butterflies was published by Kudrna (2002), and finally a butterfly field guide appeared which covered most of the West Palearctic region including all of Europe (Tshikolovets 2011).

Figure 2. 

Cumulative number of described European butterfly species per year according to current taxonomy.

The last comprehensive update of the butterfly checklist in Fauna Europaea happened 7 years ago (Karsholt and Nieukerken 2011), and the checklist presented here was first developed as an update to the online database. Unfortunately, funding for Fauna Europaea was discontinued after the initial 4-years funding period and the outdated Fauna Europaea website was only saved due to the commitment of the Natural History Museum in Berlin that set up a new one. However, its functionality is still very limited and the update process severely hampered due to shortage of funding. For this reason, we decided to publish this updated distributional checklist in order to address the need of the lepidopterological community and the public at large. It intends to cover the significant progress in butterfly systematics and faunistics, which was brought about in particular by the advancement of molecular methods.

Materials and methods

This updated checklist is based on the last version of Fauna Europaea (2.6.2). This version is almost identical to the most recent Lepidoptera update in version 2.4 (online on 28 January 2011) but includes some emendations by the staff of the Fauna Europaea office in Berlin that had not been approved by the Lepidoptera group coordinators (Erik van Nieukerken and Ole Karsholt). The geographic area covered remains the same: It includes the European mainland to the eastern slopes of the Ural mountains, plus the Macaronesian islands (excluding the Cape Verde Islands) and Cyprus, with the Caucasus and western Kazakhstan excluded (Fig. 3). Included are the British Isles and all Mediterranean islands under European administration, as well as the Greek offshore islands along the Turkish coastline. Iceland has no native butterfly species. Distributional information is based on political units at country level as in Fauna Europaea, following the ISO-3166 code. However, with the exception of the Macaronesian Islands, the additional regional splits of several countries in Fauna Europaea (mainly for Russia and some island territories) were not adopted.

Figure 3. 

Boundaries of Europe according to Fauna Europaea (from de Jong et al. 2014).

The following categories are used to explain the distribution:

A Absent (never recorded in the respective country or island group or only doubtful records)

P Present (native or well-established populations, including alien species such as the South African Cacyreus marshalli)

P? Possibly present (recorded but continued presence doubtful; usually these are species with range limits near the border of the respective country)

M Regular migrant (species which has no permanent populations, e.g., because it cannot overwinter, but is observed almost every year; included are extinct species if they are still observed as regular migrants)

I Irregular vagrant (irregular vagrants or introductions which do not reproduce or only irregularly, including temporary or recently established populations)

Ex Regionally extinct (native species which have become extinct, even though vagrants might be seen occasionally)

It should be noted that the “Extinct” category is used in a rather strict sense, in line with the IUCN Guidelines which demand that exhaustive surveys have been undertaken to prove that ‘there is no reasonable doubt that the last individual has died’. In some cases, this has led to species being recorded as “Present”, even though they are most probably extinct, e.g., Colias myrmidone in Austria (no proof for more than 25 years; H. Höttinger, pers. comm.). In addition, some of the national Red List Assessments are already outdated, even though attempts have been made to update those. An example for an update is the status of the Madeiran endemic Pieris wollastoni, whose last reliable record is from 1986. It was classified as “Critically Endangered (Possibly Extinct)” in its last Red List assessment (Van Swaay et al. 2010), but is now classified as “Extinct”, because extensive surveys in recent years have failed to prove its continued presence. This is the only European butterfly species which is known to have become globally extinct in historical times.

According to the concept of Fauna Europaea, changes were only carried out if supported by newly published research. This restriction helps to stabilize nomenclature, but can also lead to inconsistent results, e.g., due to the retention of some weakly differentiated taxa, whose species status is questionable, but for which no new published evidence is available. Potential examples in our list are Lysandra caelestissima (Verity, 1921), Polyommatus nephohiptamenos (Brown & Coutsis, 1978), Hipparchia neapolitana (Stauder, 1921), Hipparchia sbordonii Kudrna, 1984, Satyrus virbius Herrich-Schäffer, 1844, and Pieris balcana Lorković, 1969.

The main criterion whether to include or exclude a species taxon based on new (and possibly contradictory) publications was evidence for species status from at least two character sets, e.g., mitochondrial as well as nuclear DNA, or differences in morphology and karyology.

Nomenclatural changes are annotated with reference to the sources and strictly follow the last (fourth) edition of the International Code of Zoological Nomenclature (ICZN 1999). This includes the controversial article 34.2, which mandates that »the ending of a Latin or Latinised adjectival or participial species-group name must agree in gender with the generic name with which it is at any time combined«. Due to its linguistic complexity, this rule has led to many wrong or ambiguous decisions and causes additional instability of nomenclature each time a species name is transferred to another genus. Therefore a majority of lepidopterists, including the group editors of Fauna Europaea, have decided to ignore this rule and use the original spelling instead (de Jong et al. 2014). Difficulties with the gender agreement rule in Lepidoptera are as old as binominal nomenclature, because there is not even an agreement about the gender of the genus Papilio. Therefore Carl von Linné used nouns as species names and avoided the use of adjectives (Welter-Schultes 2013). However, for easy reference to Fauna Europaea and other databases, we also list the original ending and compiled a comprehensive list of original combinations, using various sources such as the LepIndex (Beccaloni et al. 2003), PESI (2018), FUNET (Savela 2018) and Tshikolovets (2011). In case of doubts or discrepancies, the original publications were checked as well.

In a few cases, necessary changes due to new nomenclatural findings have not been carried out yet, because they would result in the replacement of a well-established name by an (almost) unknown synonym. Such cases should be referred to the International Commission on Zoological Nomenclature for ruling, and changes implemented only after a decision has been made by the Commission. One such case is the well-established name Parnassius phoebus, which has turned out to represent another Asian Parnassius species which is currently known as Parnassius ariadne (Lederer, 1853) (see Hanus and Thèye 2010) and would thus need to be replaced. After the first attempt to preserve this name (Balletto and Bonelli 2014) failed (ICZN 2017), a second proposal has recently been submitted to the Commission (Lukhtanov et al. in press). According to article 82.1 of the code, prevailing usage has to be maintained until the case has been decided by the Commission.

An exceptional case which would cause a large number of changes in the names of Lepidoptera are many of the names published by [Denis & Schiffermüller] (1775) which are lacking a sufficient description, but have already been used for a very long time. In accordance with the opinion of the Fauna Europaea editorial team, we have not replaced these names. The effect on butterfly taxonomy would be rather marginal, however, because only one butterfly species would have to change its name (Nymphalis vaualbum to Nymphalis l-album (Esper, 1781)) and five others only their authorship, see Kudrna and Belicek (2005). We are looking forward to a decision of the ICZN to solve this matter (see Kudrna 2015).

Another case concerns the genus name Muschampia Tutt, 1906 (type species: Papilio proto Ochsenheimer, 1808; currently known as Muschampia proto (Ochsenheimer, 1808)), which appears to be a subjective synonym of the genus name Sloperia Tutt, 1906 (type species: Hesperia poggei Lederer, 1858; currently known as Muschampia poggei (Lederer, 1858)). Both genus names were published in the same paper and Hemming (1967) was the first to note that Sloperia should have precedence over Muschampia, because Warren (1926) as the first reviser chose Sloperia. However, the name Muschampia has remained in prevailing use during the last 90 years and, in addition, there is evidence from molecular data (Wiemers et al. unpublished) that the current classification of the species presently placed in the genera Carcharodus and Muschampia needs to be substantially revised. However, molecular data are still missing for most of the (mainly Asian) species currently placed in Muschampia, and therefore we suggest to postpone a rearrangement until better data become available.

Finally, one of us (GL) discovered that Polyommatus ottomanus Lefèbvre was published in 1831 (and not in 1830) and therefore has to be regarded as a subjective junior synonym of Lycaena legeri Freyer, 1830. This would mean that the well-established name of the species currently known as Lycaena ottomana (Lefèbvre, [1831]) would need to be changed to a name which has not been used for this species during the past century. However, according to article 23.9.1 of the Code, the prevailing usage must be maintained when the senior synonym (i.e., legeri Freyer) has not been used as a valid name after 1899 (article 23.9.1.1), and the junior synonym has been used, as its presumed valid name, in at least 25 works, published by at least ten authors during the last 50 years and encompassing a span of not less than ten years (article 23.9.1.2). In our opinion, the condition of article 23.9.1.1 applies in this case, and evidence that the conditions of article 23.9.1.2 are met, are given in Appendix 1 herein. Therefore, we regard the name Lycaena legeri Freyer as invalid and qualified as a nomen oblitum and declare the name Lycaena ottomana Lefèbvre as valid and qualified as a nomen protectum, which has precedence over the former as long as both names are thought to represent subjective synonyms.

Results and discussion

The updated species list of European butterflies includes 496 species, which belong to 110 genera in 21 subfamilies and six families (Tables 2 and 4; Fig. 4). A list of main authors with some additional data is given in Table 5. An electronic version of the checklist that includes a country-based distributional checklist is found in Suppl. material 1, Suppl. material 2.

Updated checklist of the butterflies of Europe.

Taxon Original combination Notes
Papilionidae
Papilioninae
Iphiclides podalirius (Linnaeus, 1758) Papilio podalirius
Iphiclides feisthamelii (Duponchel, 1832) Papilio feisthamelii 1
Papilio alexanor Esper, 1800 Papilio alexanor
Papilio machaon Linnaeus, 1758 Papilio machaon
Papilio hospiton Gené, 1839 Papilio hospiton 2
Parnassiinae
Parnassius mnemosyne (Linnaeus, 1758) Papilio mnemosyne
Parnassius phoebus (Fabricius, 1793) Papilio phoebus
Parnassius apollo (Linnaeus, 1758) Papilio apollo
Archon apollinus (Herbst, 1798) Papilio apollinus
Zerynthia cerisy (Godart, [1824]) Thais cerisy
Zerynthia cretica (Rebel, 1904) Thais cerisyi cretica
Zerynthia caucasica (Lederer, 1864) Thais cerisyi caucasica
Zerynthia rumina (Linnaeus, 1758) Papilio rumina
Zerynthia polyxena ([Denis & Schiffermüller], 1775) Papilio polyxena
Zerynthia cassandra (Geyer, [1828]) Papilio cassandra 3
Hesperiidae
Heteropterinae
Heteropterus morpheus (Pallas, 1771) Papilio morpheus
Carterocephalus silvicola (Meigen, 1829) Hesperia silvicola
Carterocephalus palaemon (Pallas, 1771) Papilio palaemon
Hesperiinae
Pelopidas thrax (Hübner, [1821]) Gegenes thrax
Borbo borbonica (Boisduval, 1833) Hesperia borbonica
Gegenes pumilio (Hoffmansegg, 1804) Papilio pumilio
Gegenes nostrodamus (Fabricius, 1793) Hesperia nostrodamus
Ochlodes sylvanus (Esper, 1777) Papilio sylvanus
Hesperia comma (Linnaeus, 1758) Papilio comma
Thymelicus christi Rebel, 1894 Thymelicus christi
Thymelicus acteon (Rottemburg, 1775) Papilio acteon
Thymelicus hyrax (Lederer, 1861) Hesperia hyrax
Thymelicus sylvestris (Poda, 1761) Papilio sylvestris
Thymelicus lineola (Ochsenheimer, 1808) Papilio lineola
Pyrginae
Spialia phlomidis (Herrich-Schäffer, 1845) Hesperia phlomidis
Spialia sertorius (Hoffmansegg, 1804) Hesperia sertorius
Spialia therapne (Rambur, 1832) Hesperia therapne
Spialia rosae Hernández-Roldán, Dapporto, Dincă, Vicente & Vila, 2016 Spialia rosae 4
Spialia orbifer (Hübner, [1823]) Papilio orbifer
Carcharodus tripolinus (Verity, 1925) Erynnis alceae tripolina 5
Carcharodus alceae (Esper, 1780) Papilio alceae
Muschampia cribrellum (Eversmann, 1841) Hesperia cribrellum
Muschampia tessellum (Hübner, [1803]) Papilio tessellum
Muschampia proto (Ochsenheimer, 1808) Papilio proto
Carcharodus lavatherae (Esper, 1783) Papilio lavatherae
Carcharodus orientalis Reverdin, 1913 Carcharodus orientalis
Carcharodus floccifera (Zeller, 1847) Hesperia floccifera
Carcharodus stauderi Reverdin, 1913 Carcharodus stauderi
Carcharodus baeticus (Rambur, 1839) Spilothyrus baeticus
Erynnis tages (Linnaeus, 1758) Papilio tages
Erynnis marloyi (Boisduval, 1834) Thanaos marloyi
Pyrgus malvoides (Elwes & Edwards, 1897) Hesperia malvoides
Pyrgus malvae (Linnaeus, 1758) Papilio malvae
Pyrgus carthami (Hübner, [1813]) Papilio carthami
Pyrgus sidae (Esper, 1784) Papilio sidae
Pyrgus centaureae (Rambur, 1839) Hesperia centaureae
Pyrgus cacaliae (Rambur, 1839) Hesperia cacaliae
Pyrgus andromedae (Wallengren, 1853) Syrichtus andromedae
Pyrgus serratulae (Rambur, 1839) Hesperia serratulae
Pyrgus armoricanus (Oberthür, 1910) Syrichthus armoricanus
Pyrgus alveus (Hübner, [1803]) Papilio alveus
Pyrgus warrenensis (Verity, 1928) Hesperia warrenensis
Pyrgus foulquieri (Oberthür, 1910) Syrichthus alveus foulquieri 6
Pyrgus onopordi (Rambur, 1839) Hesperia onopordi
Pyrgus carlinae (Rambur, 1839) Hesperia carlinae
Pyrgus cirsii (Rambur, 1839) Hesperia cirsii
Pyrgus cinarae (Rambur, 1839) Hesperia cinarae
Pieridae
Dismorphiinae
Leptidea duponcheli (Staudinger, 1871) Leucophasia duponcheli
Leptidea morsei (Fenton, 1882) Leptosia morsei
Leptidea juvernica Williams, 1946 Leptidea sinapis juvernica 7
Leptidea sinapis (Linnaeus, 1758) Papilio sinapis
Leptidea reali Reissinger, 1990 Leptidea sinapis reali
Coliadinae
Gonepteryx rhamni (Linnaeus, 1758) Papilio rhamni
Gonepteryx cleobule (Hübner, [1831]) Anteos cleobule 8
Gonepteryx cleopatra (Linnaeus, 1767) Papilio cleopatra
Gonepteryx maderensis C. Felder, 1862 Gonopteryx cleopatra maderensis
Gonepteryx farinosa (Zeller, 1847) Rhodocera farinosa
Catopsilia florella (Fabricius, 1775) Papilio florella
Colias hyale (Linnaeus, 1758) Papilio hyale
Colias alfacariensis Ribbe, 1905 Colias hyale alfacariensis
Colias phicomone (Esper, [1780]) Papilio phicomone
Colias aurorina Herrich-Schäffer, 1850 Colias aurorina
Colias chrysotheme (Esper, [1781]) Papilio chrysotheme
Colias erate (Esper, [1805]) Papilio erate
Colias crocea (Geoffroy, 1785) Papilio croceus 5, 9
Colias myrmidone (Esper, [1781]) Papilio myrmidone
Colias caucasica Staudinger, 1871 Colias myrmidone caucasica
Colias palaeno (Linnaeus, [1760]) Papilio palaeno 10
Colias tyche (Böber, 1812) Papilio tyche
Colias hecla Lefèbvre, 1836 Colias hecla
Pierinae
Colotis evagore (Klug, 1829) Pontia evagore
Aporia crataegi (Linnaeus, 1758) Papilio crataegi
Pontia chloridice (Hübner, [1813]) Papilio chloridice
Pontia callidice (Hübner, [1800]) Papilio callidice
Pontia edusa (Fabricius, 1777) Papilio edusa
Pontia daplidice (Linnaeus, 1758) Papilio daplidice
Pieris krueperi Staudinger, 1860 Pieris krueperi
Pieris brassicae (Linnaeus, 1758) Papilio brassicae
Pieris wollastoni (Butler, 1886) Ganoris wollastoni
Pieris cheiranthi (Hübner, [1808]) Papilio cheiranthi
Pieris rapae (Linnaeus, 1758) Papilio rapae
Pieris mannii (Mayer, 1851) Pontia mannii
Pieris ergane (Geyer, [1828]) Papilio ergane
Pieris bryoniae (Hübner, [1806]) Papilio bryoniae
Pieris napi (Linnaeus, 1758) Papilio napi
Pieris balcana Lorković, [1969] Pieris balcana 11
Euchloe tagis (Hübner, [1804]) Papilio tagis
Euchloe eversi Stamm, 1963 Euchloe belemia eversi
Euchloe grancanariensis Acosta, 2008 Euchloe belemia grancanariensis
Euchloe hesperidum Rothschild, 1913 Euchloe belemia hesperidum
Euchloe belemia (Esper, 1800) Papilio belemia
Euchloe insularis (Staudinger, 1861) Anthocharis tagis insularis
Euchloe crameri Butler, 1869 Euchloe crameri
Euchloe simplonia (Freyer, 1829) Pontia simplonia
Euchloe ausonia (Hübner, [1804]) Papilio ausonia
Euchloe charlonia (Donzel, 1842) Anthocharis charlonia
Euchloe penia (Freyer, 1851) Pontia penia
Euchloe bazae Fabiano, 1993 Euchloe charlonia bazae
Zegris pyrothoe (Eversmann, 1832) Pontia pyrothoe
Zegris eupheme (Esper, [1804]) Papilio eupheme
Anthocharis euphenoides Staudinger, 1869 Anthocharis euphenoides
Anthocharis cardamines (Linnaeus, 1758) Papilio cardamines
Anthocharis gruneri Herrich-Schäffer, 1851 Anthocharis gruneri
Anthocharis damone Boisduval, 1836 Anthocharis damone
Riodinidae
Nemeobiinae
Hamearis lucina (Linnaeus, 1758) Papilio lucina
Lycaenidae
Lycaeninae
Lycaena dimorpha (Staudinger, 1881) Polyommatus dimorphus 5, 12
Lycaena helle ([Denis & Schiffermüller], 1775) Papilio helle
Lycaena alciphron (Rottemburg, 1775) Papilio alciphron
Lycaena thetis Klug, 1834 Lycaena thetis
Lycaena thersamon (Esper, 1784) Papilio thersamon
Lycaena dispar ([Haworth], 1802) Papilio dispar
Lycaena hippothoe (Linnaeus, [1760]) Papilio hippothoe 10
Lycaena candens (Herrich-Schäffer, 1844) Polyommatus candens
Lycaena ottomana (Lefèbvre, [1831]) Polyommatus ottomanus 5, 13
Lycaena bleusei (Oberthür, 1884) Polyommatus xanthe f. bleusei
Lycaena phlaeas (Linnaeus, [1760]) Papilio phlaeas 10
Lycaena virgaureae (Linnaeus, 1758) Papilio virgaureae
Lycaena tityrus (Poda, 1761) Papilio tityrus
Aphnaeinae
Cigaritis acamas (Klug, 1834) Lycaena acamas 14
Theclinae
Thecla betulae (Linnaeus, 1758) Papilio betulae
Favonius quercus (Linnaeus, 1758) Papilio quercus
Laeosopis roboris (Esper, [1793]) Papilio roboris 15
Tomares ballus (Fabricius, 1787) Papilio ballus
Tomares nogelii (Herrich-Schäffer, 1851) Thecla nogelii
Tomares callimachus (Eversmann, 1848) Lycaena callimachus
Callophrys avis Chapman, 1909 Callophrys avis
Callophrys suaveola (Staudinger, 1881) Thecla suaveola
Callophrys rubi (Linnaeus, 1758) Papilio rubi
Callophrys chalybeitincta Sovinsky, 1905 Callophrys rubi chalybeitincta
Neolycaena rhymnus (Eversmann, 1832) Lycaena rhymnus
Satyrium pruni (Linnaeus, 1758) Papilio pruni
Satyrium ilicis (Esper, 1779) Papilio ilicis
Satyrium esculi (Hübner, [1804]) Papilio esculi
Satyrium ledereri (Boisduval, 1848) Lycaena ledereri
Satyrium w-album (Knoch, 1782) Papilio w-album
Satyrium spini ([Denis & Schiffermüller], 1775) Papilio spini
Satyrium acaciae (Fabricius, 1787) Papilio acaciae
Polyommatinae
Leptotes pirithous (Linnaeus, 1767) Papilio pirithous
Cyclyrius webbianus (Brullé, 1839) Polyommatus webbianus
Azanus ubaldus (Stoll, 1782) Papilio ubaldus
Azanus jesous (Guérin-Méneville, 1849) Polyommatus jesous
Lampides boeticus (Linnaeus, 1767) Papilio boeticus
Cacyreus marshalli Butler, 1898 Cacyreus marshalli
Celastrina argiolus (Linnaeus, 1758) Papilio argiolus
Tarucus theophrastus (Fabricius, 1793) Hesperia theophrastus
Tarucus balkanicus (Freyer, 1844) Lycaena balkanica 5
Phengaris alcon ([Denis & Schiffermüller], 1775) Papilio alcon
Phengaris arion (Linnaeus, 1758) Papilio arion
Phengaris teleius (Bergsträsser, 1779) Papilio teleius
Phengaris nausithous (Bergsträsser, 1779) Papilio nausithous
Turanana taygetica (Rebel, 1902) Lycaena panagaea taygetica
Pseudophilotes bavius (Eversmann, 1832) Lycaena bavius
Pseudophilotes barbagiae De Prins & van der Poorten, 1982 Pseudophilotes barbagiae
Pseudophilotes abencerragus (Pierret, 1837) Argus abencerragus
Pseudophilotes panoptes (Hübner, [1813]) Papilio panoptes
Pseudophilotes vicrama (Moore, 1865) Polyommatus vicrama
Pseudophilotes baton (Bergsträsser, 1779) Papilio baton
Scolitantides orion (Pallas, 1771) Papilio orion
Praephilotes anthracias (Christoph, 1877) Lycaena anthracias
Iolana iolas (Ochsenheimer, 1816) Lycaena iolas
Iolana debilitata (Schultz, 1905) Lycaena jolas var. debilitata 16
Glaucopsyche melanops (Boisduval, 1828) Polyommatus melanops
Glaucopsyche paphos Chapman, 1920 Glaucopsyche paphos
Glaucopsyche alexis (Poda, 1761) Papilio alexis
Zizeeria knysna (Trimen, 1862) Lycaena knysna
Zizeeria karsandra (Moore, 1865) Polyommatus karsandra
Tongeia fischeri (Eversmann, 1843) Lycaena fischeri
Cupido argiades (Pallas, 1771) Papilio argiades
Cupido decoloratus (Staudinger, 1886) Lycaena argiades decolorata 5
Cupido alcetas (Hoffmansegg, 1804) Papilio alcetas
Cupido osiris (Meigen, 1829) Polyommatus osiris
Cupido minimus (Fuessly, 1775) Papilio minimus
Cupido lorquinii (Herrich-Schäffer, 1850) Lycaena lorquinii 17
Luthrodes galba (Lederer, 1855) Lycaena galba 18
Freyeria trochylus (Freyer, 1844) Lycaena trochylus 18,19
Plebejus argus (Linnaeus, 1758) Papilio argus
Plebejus idas (Linnaeus, [1760]) Papilio idas 10
Plebejus bellieri (Oberthür, 1910) Lycaena bellieri
Plebejus argyrognomon (Bergsträsser, 1779) Papilio argyrognomon
Agriades orbitulus (Prunner, 1798) Papilio orbitulus 18
Agriades optilete (Knoch, 1781) Papilio optilete 18
Agriades pyrenaicus (Boisduval, 1840) Lycaena orbitulus var. pyrenaica 5, 18
Agriades dardanus (Freyer, 1843) Lycaena dardanus 18
Agriades zullichi Hemming, 1933 Agriades zullichi 18
Agriades glandon (Prunner, 1798) Papilio glandon 18
Agriades aquilo (Boisduval, 1832) Argus aquilo 18
Plebejidea loewii (Zeller, 1847) Lycaena loewii 18
Eumedonia eumedon (Esper, 1780) Papilio eumedon 18
Kretania psylorita (Freyer, 1845) Lycaena psylorita 18
Kretania hesperica (Rambur, 1839) Polyommatus hespericus 5, 18
Kretania eurypilus (Freyer, 1851) Lycaena eurypilus 18
Kretania trappi (Verity, 1927) Lycaena trappi 18
Kretania sephirus (Frivaldszky, 1835) Lycaena sephirus 18
Kretania pylaon (Fischer, 1832) Lycaena pylaon 18
Cyaniris semiargus (Rottemburg, 1775) Papilio semiargus
Glabroculus cyane (Eversmann, 1837) Lycaena cyane 18
Aricia morronensis (Ribbe, 1910) Lycaena idas morronensis
Aricia anteros (Freyer, 1838) Lycaena anteros
Aricia cramera (Eschscholtz, 1821) Lycaena cramera
Aricia nicias (Meigen, 1829) Polyommatus nicias 20
Aricia artaxerxes (Fabricius, 1793) Hesperia artaxerxes
Aricia montensis Verity, 1928 Aricia medon montensis
Aricia agestis ([Denis & Schiffermüller], 1775) Papilio agestis
Neolysandra coelestina (Eversmann, 1843) Lycaena coelestina 18
Lysandra hispana (Herrich-Schäffer, 1851) Lycaena coridon var. hispana 18
Lysandra corydonius (Herrich-Schäffer, 1852) Lycaena coridon corydonius 18
Lysandra bellargus (Rottemburg, 1775) Papilio bellargus 18
Lysandra coridon (Poda, 1761) Papilio coridon 18
Lysandra caelestissima (Verity, 1921) Agriades coridon caelestissima 18
Lysandra albicans (Gerhard, 1851) Lycaena coridon var. albicans 18
Polyommatus escheri (Hübner, [1823]) Papilio escheri
Polyommatus thersites (Cantener, 1835) Argus thersites
Polyommatus daphnis ([Denis & Schiffermüller], 1775) Papilio daphnis
Polyommatus amandus (Schneider, 1792) Papilio amandus
Polyommatus golgus (Hübner, [1813]) Papilio golgus
Polyommatus nivescens (Keferstein, 1851) Lycaena dorylas var. nivescens
Polyommatus dorylas ([Denis & Schiffermüller], 1775) Papilio dorylas
Polyommatus celina (Austaut, 1879) Lycaena celina 21
Polyommatus icarus (Rottemburg, 1775) Papilio icarus
Polyommatus eros (Ochsenheimer, 1808) Papilio eros
Polyommatus damon ([Denis & Schiffermüller], 1775) Papilio damon
Polyommatus damone (Eversmann, 1841) Lycaena damone
Polyommatus damocles (Herrich-Schäffer, 1844) Lycaena damocles
Polyommatus admetus (Esper, 1783) Papilio admetus
Polyommatus ripartii (Freyer, 1830) Lycaena ripartii
Polyommatus nephohiptamenos (Brown & Coutsis, 1978) Agrodiaetus nephohiptamenos
Polyommatus iphigenia (Herrich-Schäffer, 1847) Lycaena iphigenia
Polyommatus violetae (Gómez-Bustillo, Expósito & Martínez, 1979) Agrodiaetus violetae
Polyommatus fulgens (Sagarra, 1925) Hirsutina dolus r. fulgens 22
Polyommatus fabressei (Oberthür, 1910) Lycaena rippertii r. fabressei
Polyommatus dolus (Hübner, [1823]) Papilio dolus
Polyommatus humedasae (Toso & Balletto, 1976) Agrodiaetus humedasae
Polyommatus timfristos Lukhtanov, Vishnevskaya & Shapoval, 2016 Polyommatus timfristos 23
Polyommatus orphicus Kolev, 2005 Polyommatus orphicus
Polyommatus aroaniensis (Brown, 1976) Agrodiaetus alcestis aroaniensis
Nymphalidae
Limenitidinae
Neptis sappho (Pallas, 1771) Papilio sappho
Neptis rivularis (Scopoli, 1763) Papilio rivularis
Limenitis reducta Staudinger, 1901 Limenitis camilla reducta
Limenitis populi (Linnaeus, 1758) Papilio populi
Limenitis camilla (Linnaeus, 1764) Papilio camilla
Heliconiinae
Issoria lathonia (Linnaeus, 1758) Papilio lathonia
Issoria eugenia (Eversmann, 1847) Argynnis eugenia
Brenthis hecate ([Denis & Schiffermüller], 1775) Papilio hecate
Brenthis ino (Rottemburg, 1775) Papilio ino
Brenthis daphne ([Denis & Schiffermüller], 1775) Papilio daphne
Argynnis paphia (Linnaeus, 1758) Papilio paphia
Argynnis pandora ([Denis & Schiffermüller], 1775) Papilio pandora
Argynnis laodice (Pallas, 1771) Papilio laodice
Speyeria aglaja (Linnaeus, 1758) Papilio aglaja 24
Fabriciana elisa (Godart, 1823) Argynnis elisa 24
Fabriciana niobe (Linnaeus, 1758) Papilio niobe 24
Fabriciana adippe ([Denis & Schiffermüller], 1775) Papilio adippe 24
Boloria eunomia (Esper, 1800) Papilio eunomia 25
Boloria graeca (Staudinger, 1870) Argynnis pales graeca
Boloria pales ([Denis & Schiffermüller], 1775) Papilio pales
Boloria alaskensis (Holland, 1900) Argynnis alaskensis
Boloria napaea (Hoffmansegg, 1804) Papilio napaea
Boloria aquilonaris (Stichel, 1908) Argynnis aquilonaris
Boloria tritonia (Böber, 1812) Papilio tritonia
Boloria polaris (Boisduval, 1828) Argynnis polaris
Boloria thore (Hübner, [1804]) Papilio thore 26
Boloria selene ([Denis & Schiffermüller], 1775) Papilio selene
Boloria euphrosyne (Linnaeus, 1758) Papilio euphrosyne
Boloria dia (Linnaeus, 1767) Papilio dia
Boloria improba (Butler, 1877) Argynnis improba
Boloria frigga (Thunberg, 1791) Papilio frigga 27
Boloria freija (Thunberg, 1791) Papilio freija 27
Boloria selenis (Eversmann, 1837) Argynnis selenis
Boloria oscarus (Eversmann, 1844) Argynnis oscarus
Boloria titania (Esper, [1793]) Papilio titania
Boloria chariclea (Schneider, 1794) Papilio chariclea
Boloria angarensis (Erschoff, 1870) Argynnis angarensis
Apaturinae
Apatura iris (Linnaeus, 1758) Papilio iris
Apatura metis Freyer, 1829 Apatura metis
Apatura ilia ([Denis & Schiffermüller], 1775) Papilio ilia
Nymphalinae
Araschnia levana (Linnaeus, 1758) Papilio levana
Vanessa virginiensis (Drury, 1773) Papilio cardui virginiensis
Vanessa cardui (Linnaeus, 1758) Papilio cardui
Vanessa vulcania Godart, 1819 Vanessa vulcania
Vanessa atalanta (Linnaeus, 1758) Papilio atalanta
Aglais io (Linnaeus, 1758) Papilio io
Aglais urticae (Linnaeus, 1758) Papilio urticae
Aglais ichnusa (Hübner, [1824]) Papilio ichnusa 28
Polygonia egea (Cramer, 1775) Papilio egea
Polygonia c-album (Linnaeus, 1758) Papilio c-album
Nymphalis vaualbum ([Denis & Schiffermüller], 1775) Papilio vau album
Nymphalis polychloros (Linnaeus, 1758) Papilio polychloros
Nymphalis xanthomelas ([Denis & Schiffermüller], 1775) Papilio xanthomelas
Nymphalis antiopa (Linnaeus, 1758) Papilio antiopa
Hypolimnas misippus (Linnaeus, 1764) Papilio misippus
Euphydryas desfontainii (Godart, 1819) Papilio desfontainii
Euphydryas aurinia (Rottemburg, 1775) Papilio aurinia
Euphydryas cynthia ([Denis & Schiffermüller], 1775) Papilio cynthia
Euphydryas iduna (Dalman, 1816) Melitaea iduna
Euphydryas maturna (Linnaeus, 1758) Papilio maturna
Euphydryas intermedia (Ménétriés, 1859) Melitaea maturna intermedia
Melitaea trivia ([Denis & Schiffermüller], 1775) Papilio trivia
Melitaea didyma (Esper, 1778) Papilio didyma
Melitaea arduinna (Esper, 1783) Papilio arduinna
Melitaea aetherie (Hübner, [1826]) Papilio aetherie
Melitaea phoebe ([Denis & Schiffermüller], 1775) Papilio phoebe
Melitaea ornata Christoph, 1893 Melitaea phoebe ornata 29
Melitaea cinxia (Linnaeus, 1758) Papilio cinxia
Melitaea diamina (Lang, 1789) Papilio diamina
Melitaea celadussa Fruhstorfer, 1910 Melitaea athalia celadussa 30
Melitaea deione (Geyer, [1832]) Papilio deione
Melitaea britomartis Assmann, 1847 Melitaea britomartis
Melitaea athalia (Rottemburg, 1775) Papilio athalia
Melitaea varia Herrich-Schäffer, 1851 Melitaea varia 31
Melitaea parthenoides Keferstein, 1851 Melitaea athalia parthenoides
Melitaea aurelia Nickerl, 1850 Melitaea aurelia
Melitaea asteria Freyer, 1828 Melitaea asteria
Libytheinae
Libythea celtis (Laicharting, 1782) Papilio celtis
Danainae
Danaus plexippus (Linnaeus, 1758) Papilio plexippus
Danaus chrysippus (Linnaeus, 1758) Papilio chrysippus
Charaxinae
Charaxes jasius (Linnaeus, 1767) Papilio jasius
Satyrinae
Coenonympha phryne (Pallas, 1771) Papilio phryne
Coenonympha oedippus (Fabricius, 1787) Papilio oedippus
Coenonympha dorus (Esper, 1782) Papilio dorus
Coenonympha thyrsis (Freyer, 1845) Hipparchia thyrsis
Coenonympha pamphilus (Linnaeus, 1758) Papilio pamphilus
Coenonympha tullia (Müller, 1764) Papilio tullia
Coenonympha rhodopensis Elwes, 1900 Coenonympha tiphon rhodopensis
Coenonympha amaryllis (Stoll, 1782) Papilio amaryllis
Coenonympha glycerion (Borkhausen, 1788) Papilio glycerion
Coenonympha corinna (Hübner, [1804]) Papilio corinna
Coenonympha leander (Esper, 1784) Papilio leander
Coenonympha hero (Linnaeus, [1760]) Papilio hero 10
Coenonympha gardetta (Prunner, 1798) Papilio gardetta
Coenonympha orientalis Rebel, 1909 Coenonympha arcania var. orientalis 32
Coenonympha arcania (Linnaeus, [1760]) Papilio arcania 10
Kirinia roxelana (Cramer, 1777) Papilio roxelana
Kirinia climene (Esper, 1783) Papilio climene
Lopinga achine (Scopoli, 1763) Papilio achine
Pararge xiphia (Fabricius, 1775) Papilio xiphia
Pararge xiphioides Staudinger, 1871 Pararge xiphia xiphioides
Pararge aegeria (Linnaeus, 1758) Papilio aegeria
Lasiommata maera (Linnaeus, 1758) Papilio maera
Lasiommata deidamia (Eversmann, 1851) Hipparchia deidamia
Lasiommata petropolitana (Fabricius, 1787) Papilio maera petropolitana
Lasiommata paramegaera (Hübner, [1824]) Papilio paramegaera
Lasiommata megera (Linnaeus, 1767) Papilio megera
Melanargia russiae (Esper, 1783) Papilio russiae
Melanargia larissa (Geyer, [1828]) Papilio larissa
Melanargia lachesis (Hübner, 1790) Papilio lachesis
Melanargia galathea (Linnaeus, 1758) Papilio galathea
Melanargia ines (Hoffmansegg, 1804) Papilio ines
Melanargia arge (Sulzer, 1776) Papilio arge
Melanargia pherusa (Boisduval, 1833) Arge pherusa
Melanargia occitanica (Esper, [1793]) Papilio arge occitanica
Hipparchia fatua Freyer, 1843 Hipparchia fatua 33
Hipparchia statilinus (Hufnagel, 1766) Papilio statilinus
Hipparchia tilosi Manil, 1984 Hipparchia wyssii tilosi
Hipparchia bacchus (Higgins, 1967) Pseudotergumia wyssii bacchus
Hipparchia wyssii (Christ, 1889) Satyrus fidia wyssii
Hipparchia tamadabae Owen & Smith, 1992 Hipparchia wyssi tamadabae
Hipparchia gomera (Higgins, 1967) Pseudotergumia wyssii gomera
Hipparchia fidia (Linnaeus, 1767) Papilio fidia
Hipparchia neomiris (Godart, 1823) Satyrus neomiris 34
Hipparchia autonoe (Esper, 1783) Papilio autonoe
Hipparchia hermione (Linnaeus, 1764) Papilio hermione
Hipparchia syriaca (Staudinger, 1871) Satyrus hermione syriaca
Hipparchia fagi (Scopoli, 1763) Papilio fagi
Hipparchia mersina (Staudinger, 1871) Satyrus semele mersina
Hipparchia miguelensis (Le Cerf, 1935) Satyrus azorinus miguelensis
Hipparchia azorina (Strecker, 1899) Satyrus azorinus 5, 35
Hipparchia senthes (Fruhstorfer, 1908) Eumenis semele senthes
Hipparchia maderensis (Bethune-Baker, 1891) Satyrus semele maderensis
Hipparchia semele (Linnaeus, 1758) Papilio semele
Hipparchia blachieri (Fruhstorfer, 1908) Eumenis semele blachieri
Hipparchia aristaeus (Bonelli, 1826) Papilio aristaeus
Hipparchia volgensis (Mazokhin-Porshnyakov, 1952) Satyrus semele volgensis
Hipparchia neapolitana (Stauder, 1921) Satyrus neapolitana
Hipparchia leighebi Kudrna, 1976 Hipparchia semele leighebi
Hipparchia pellucida (Stauder, 1924) Satyrus semele pellucida 36
Hipparchia sbordonii Kudrna, 1984 Hipparchia sbordonii
Hipparchia cypriensis (Holik, 1949) Satyrus semele cypriensis
Hipparchia cretica (Rebel, 1916) Satyrus semele cretica
Hipparchia christenseni Kudrna, 1977 Hipparchia christenseni
Minois dryas (Scopoli, 1763) Papilio dryas
Brintesia circe (Fabricius, 1775) Papilio circe
Arethusana arethusa ([Denis & Schiffermüller], 1775) Papilio arethusa
Oeneis tarpeia (Pallas, 1771) Papilio tarpeia
Oeneis bore (Schneider, 1792) Papilio bore
Oeneis ammon Elwes, 1899 Oeneis bore var. ammon 37
Oeneis melissa (Fabricius, 1775) Papilio melissa
Oeneis magna Graeser, 1888 Oeneis jutta magna
Oeneis jutta (Hübner, [1806]) Papilio jutta
Oeneis norna (Thunberg, 1791) Papilio norna
Oeneis polixenes (Fabricius, 1775) Papilio polixenes
Oeneis glacialis (Moll, 1785) Papilio glacialis 38
Satyrus ferula (Fabricius, 1793) Papilio ferula
Satyrus virbius Herrich-Schäffer, 1844 Satyrus virbius
Satyrus actaea (Esper, 1781) Papilio actaea
Chazara briseis (Linnaeus, 1764) Papilio briseis
Chazara prieuri (Pierret, 1837) Satyrus prieuri
Chazara persephone (Hübner, [1805]) Papilio persephone
Pseudochazara geyeri (Herrich-Schäffer, 1846) Satyrus geyeri
Pseudochazara graeca (Staudinger, 1870) Satyrus pelopea graeca
Pseudochazara amymone Brown, 1976 Pseudochazara amymone
Pseudochazara anthelea (Hübner, [1824]) Papilio anthelea
Pseudochazara amalthea (Frivaldszky, 1845) Hipparchia amalthea 39
Pseudochazara williamsi (Romei, 1927) Satyrus hippolyte williamsi
Pseudochazara euxina (Kuznetsov, 1909) Hipparchia euxina
Pseudochazara mercurius (Staudinger, 1887) Satyrus mercurius 40
Pseudochazara cingovskii (Gross, 1973) Satyrus sintenisi cingovskii
Pseudochazara tisiphone Brown, [1981] Pseudochazara cingovskii tisiphone 39
Pseudochazara orestes De Prins & van der Poorten, 1981 Pseudochazara orestes
Ypthima asterope (Klug, 1832) Hipparchia asterope
Proterebia phegea (Borkhausen, 1788) Papilio phegea 41
Hyponephele huebneri Koçak, 1980 Hyponephele huebneri
Hyponephele lycaon (Kühn, 1774) Papilio lycaon
Hyponephele lupina (Costa, 1836) Satyrus lupinus 5
Aphantopus hyperantus (Linnaeus, 1758) Papilio hyperantus
Pyronia cecilia (Vallantin, 1894) Epinephele ida cecilia
Pyronia tithonus (Linnaeus, 1771) Papilio tithonus 42
Pyronia bathseba (Fabricius, 1793) Papilio bathseba
Maniola jurtina (Linnaeus, 1758) Papilio jurtina
Maniola nurag (Ghiliani, 1852) Satyrus nurag
Maniola chia Thomson, 1987 Maniola chia
Maniola megala (Oberthür, 1909) Epinephele janira megala
Maniola cypricola (Graves, 1928) Epinephele cypricola
Maniola telmessia (Zeller, 1847) Hipparchia telmessia
Maniola halicarnassus Thomson, 1990 Maniola halicarnassus
Erebia edda Ménétriés, 1851 Erebia edda
Erebia fasciata Butler, 1868 Erebia fasciata
Erebia discoidalis (Kirby, 1837) Hipparchia discoidalis
Erebia rossii (Curtis, 1835) Hipparchia rossii 43
Erebia cyclopius (Eversmann, 1844) Hipparchia cyclopius
Erebia embla (Thunberg, 1791) Papilio embla
Erebia disa (Thunberg, 1791) Papilio disa
Erebia meolans (Prunner, 1798) Papilio meolans
Erebia dabanensis Erschoff, 1872 Erebia dabanensis 44
Erebia jeniseiensis Trybom, 1877 Erebia ligea jeniseiensis
Erebia claudina (Borkhausen, 1789) Papilio claudina
Erebia manto ([Denis & Schiffermüller], 1775) Papilio manto
Erebia ottomana Herrich-Schäffer, 1847 Erebia dromus ottomana
Erebia hispania Butler, 1868 Erebia hispania
Erebia rondoui Oberthür, 1908 Erebia rondoui
Erebia callias Edwards, 1871 Erebia callias 45
Erebia tyndarus (Esper, 1781) Papilio tyndarus
Erebia cassioides (Hohenwarth, 1792) Papilio cassioides 46
Erebia nivalis Lorković & Lesse, 1954 Erebia nivalis
Erebia neleus (Freyer, 1832) Hipparchia neleus 47
Erebia calcarius Lorković, 1953 Erebia tyndarus calcarius
Erebia arvernensis Oberthür, 1908 Erebia tyndarus arvernensis 47
Erebia oeme (Hübner, [1804]) Papilio oeme
Erebia gorge (Hübner, [1804]) Papilio gorge
Erebia sthennyo Graslin, 1850 Erebia sthennyo
Erebia pandrose (Borkhausen, 1788) Papilio pandrose
Erebia eriphyle (Freyer, 1836) Hipparchia eriphyle
Erebia epistygne (Hübner, [1819]) Papilio epistygne
Erebia euryale (Esper, 1805) Papilio euryale
Erebia palarica Chapman, 1905 Erebia palarica
Erebia ligea (Linnaeus, 1758) Papilio ligea
Erebia pluto (Prunner, 1798) Papilio pluto
Erebia aethiopellus (Hoffmansegg, 1806) Papilio aethiopellus
Erebia gorgone Boisduval, 1833 Erebia gorgone
Erebia rhodopensis Nicholl, 1900 Erebia gorgone rhodopensis
Erebia mnestra (Hübner, [1804]) Papilio mnestra
Erebia albergana (Prunner, 1798) Papilio alberganus 5
Erebia sudetica Staudinger, 1861 Erebia melampus sudetica
Erebia melampus (Fuessly, 1775) Papilio melampus
Erebia triarius (Prunner, 1798) Papilio triarius
Erebia polaris Staudinger, 1861 Erebia medusa var. polaris 48
Erebia medusa ([Denis & Schiffermüller], 1775) Papilio medusa
Erebia aethiops (Esper, 1777) Papilio aethiops
Erebia pharte (Hübner, [1804]) Papilio pharte
Erebia christi Rätzer, 1890 Erebia christi
Erebia orientalis Elwes, 1900 Erebia epiphron orientalis
Erebia epiphron (Knoch, 1783) Papilio epiphron
Erebia flavofasciata Heyne, 1895 Erebia flavofasciata
Erebia montana (Prunner, 1798) Papilio montanus 5
Erebia styx (Freyer, 1834) Hipparchia styx
Erebia stiria (Godart, [1824]) Satyrus stirius 5
Erebia scipio Boisduval, 1833 Erebia scipio 49
Erebia pronoe (Esper, 1780) Papilio pronoe
Erebia melas (Herbst, 1796) Papilio melas
Erebia lefebvrei (Boisduval, 1828) Satyrus lefebvrei
Erebia zapateri Oberthür, 1875 Erebia zapateri
Erebia neoridas (Boisduval, 1828) Satyrus neoridas

Annotations to the updated checklist of the butterflies of Europe.

1 Iphiclides feisthamelii is considered a separate species based on differences in adult morphology (Coutsis and van Oorschot 2011, Lafranchis et al. 2015) and nuclear genetic markers (Wiemers and Gottsberger 2010; Dincă et al. 2015), despite very local hybridisation along the contact zone in southern France (Lafranchis et al. 2015) and extensive mitochondrial introgression in the Iberian Peninsula (Wiemers and Gottsberger 2010; Dincă et al. 2015). Its distribution includes the SW part of France, the Iberian Peninsula, and northern Africa.
2 Author of the name is Giuseppe Gené (1800–1847), not Achille Guenée.
3 Dapporto (2009) has shown that Zerynthia cassandra from peninsular Italy is a separate species based on differences in genital morphology. This was further confirmed by molecular studies (Zinetti et al. 2013).
4 Spialia rosae has been recognised as a separate species endemic to mountains of Spain based on differences in ecology and evidence from molecular studies (mitochondrial DNA, chemical profiles) (Hernández-Roldán et al. 2016, 2018). The species has already been included in Fauna Europaea (2018).
5 Gender agreement changes were applied consistently in accordance with Art. 31.2 and Art. 34.2 (ICZN 1999).
6 As descriptions of both Syrichtus alveus f. foulquieri and Syrichtusalveus f. bellieri were published simultaneously (Oberthür, 1910), the name used by the first reviser (i. e. Rebel 1914), Pyrgus foulquieri, should be used in accordance with Art. 24.2.1 and Art. 24.2.2 (ICZN 1999).
7 Recent studies have shown that Leptidea reali actually comprises two species, L. reali and L. juvernica. L. reali is known from south-western Europe (Spain, S France and Italy) and is replaced by L. juvernica in the rest of the continent (Dincă et al. 2011b). L. sinapis, L. reali, and L. juvernica are reproductively isolated due to female mate choice (Dincă et al. 2013).
8 The year of the publication of the name Anteos cleobule is 1831, not 1830 (the original plate [79], published in 1824, carried no names).
9 The name Papilio croceus should be credited to Geoffroy in Fourcroy, 1785, not to Fourcroy (Ganglbauer and Heyden 1906, D'Aguilar and Raimbault 1990, Grieshuber et al. 2012).
10 The date of the publication of the names by Linnaeus in Fauna Svecica (ed. 2) is 14 November 1760, not 1761 (see Evenhuis 1997, Bousquet 2016).
11 The year of the publication of the name Pieris balcana is 1969, not 1970. The publication year of volume 21 (1–4) (1968) of Biološki glasnik [= volume 70 of Periodicum Biologorum] is printed on the cover page as “1969” and, moreover, Lorković´s personal copy held in the Croatian Natural History museum has a hand written addition of the publication year “1969” in the header of his article (Šašić, pers. comm.). Additionally, the author´s name is misspelled and should be Lorković (see also Lorković 1969).
12 According to Lvovsky and Morgun (2007) the species is present in Russia south of the Urals in the Orenburg region. The subspecies Lycaena dimorpha irghiza was originally described as a subspecies of L. japhetica (Nekrutenko 1985), but we follow the decision in the taxonomic review by Lukhtanov (2000).
13 The year of the publication of the name Polyommatus ottomanus is 1831, not 1830. Lefèbvre cited the date 1830, which corresponds to the date of submission of his article, but the issue of the journal was published in January 1831. See Lefèbvre (1831)
14 The generic names Apharitis and Spindasis were synonymised with Cigaritis due to morphological similarities (see Heath and Pringle 2011).
15 The name Papilio roboris was first published in 1793, not 1789 (Lamas 2013).
16 Iolana debilitata has been recognised as a separate species based on constant differences in adult morphology (Dumont 2004) and mitochondrial DNA – barcoding gene (Dincă et al. 2015).
17 The year of the publication of the name and plates for Cupido lorquinii is 1850, not 1847 (Hemming 1937, Heppner 1982).
18 Genus level classification in the subfamily Polyommatinae follows Talavera et al. (2013) based on molecular phylogeny. This arrangement partially concurs with differences in genital morphology (see Balletto et al. 2014, Coutsis 2017).
19 The year of the publication of the name Lycaena trochylus is 1844, not 1845 (Tremewan 1988, Olivier 2000).
20 The year of the publication of the name Polyommatus nicias is ante September 1829, not 1830 (Griffin 1931).
21 Polyommatus celina has been recognised as a separate species distributed in the Iberian Peninsula, northern Africa, Sardinia and Sicily based on molecular markers and adult morphology (Wiemers et al. 2010; Dincă et al. 2011a).
22 The author´s surname Sagarra should be without the particle “de”. It is listed as such in the members list of the Institució Catalana d'Història Natural in 1925 bulletin Vol. 5 – Num. 1. Generally, when the particle is written in lowercase, it should be treated as a suffix that goes after the first name (Welter-Schultes 2013).
23 Polyommatus timfristos is considered a separate species due to differences in haploid chromosome number compared to P. aroaniensis and mitochondrial DNA – barcoding gene (Vishnevskaya et al. 2016).
24 Genus level classification in the tribe Argynnini follows De Moya et al. (2017) based on molecular phylogenetics. It is corroborated by extensive differences in genital morphology (Simonsen 2006a, 2006b).
25 The name Papilio eunomia was first published in 1800, not 1799 (Poche, 1938).
26 The name Papilio thore was first published in 1804, not 1803 (Hemming 1937).
27 Description of Boloria freija and Boloria frigga must be credited to Thunberg, not to Becklin (Thunberg wrote Becklin's dissertation), see Karsholt and Nielsen (1986).
28 Papilio ichnusa was first described by Hübner (ante 23 December) 1824. Vanessa ichnusa Bonelli was published in February 1825 and is a junior secondary homonym and junior subjective synonym, see Hemming (1937).
29 Among the species with red headed larvae within the Melitaea phoebe species group only M. ornata is present in Europe in southeastern Russia, the Balkan Peninsula, Spain, southeastern France, and southern Italy. M. telona is limited to the Levant and M. punica to northern Africa (Toth et al. 2014).
30 Melitaea celadussa Fruhstorfer, 1910 is considered a separate species distributed in western Europe that differs in genital morphology (Higgins 1932) and molecular markers (Leneveu et al. 2009, Dincă et al. 2015) from M. athalia, with hybrids known from the contact zone (Achtelik 2006; Oorschot and Coutsis 2014). The species was referred to also as M. nevadensis Oberthür, 1904, which is a junior primary homonym of Melitaea parthenie var. nevadensis Spuler, 1901, currently regarded as a junior subjective synonym of Melitaea parthenoides Keferstein, 1851.
31 Melitaea varia was first described by Herrich-Schäffer (1851) in Systematische Bearbeitung der Schmetterlinge von Europa Vol. 6(48): 2 (Hemming 1937). Melitaea parthenie var. varia Meyer-Dür, 1852 (not 1851) is a junior primary homonym.
32 The name Coenonympha arcánia var. orientális [sic] appeared in part 4 of the ninth edition of Berge’s Schmetterlingsbuch, which was published on 22 May 1909 (Lempke 1949), not in 1910.
33 The name Hipparchia fatua was first published in 1843, not 1844 (Olivier 2000).
34 The name Satyrus neomiris was first published in 1823, not 1822. Satyrus neomiris first appeared on page 19 in Godart’s Tableau méthodique des lépidoptères..., published in 1823. The vernacular name Godart used in vol. 2 of Hist. nat. Lépid. Pap. France, pp. 88–89, pl. 11, figs. 1–2 (1822), »Satyre néomiris«, is unavailable, as it is not a scientific name.
35 The name Satyrus azorinus was first published in 1899, not 1898.
36 The name Satyrus semele pellucida was first published on 15 May 1924, not in 1923.
37 Oeneis ammon is present in Europe in the Polar Urals (Tsvetkov 2006).
38 The name Papilio glacialis was first published in 1785, not 1783.
39 Based on differentiation in mtDNA (barcodes) and differences in morphology, Pseudochazara amalthea and P. tisiphone are considered separate species from allopatric P. anthelea and P. mniszechii respectively (Verovnik and Wiemers 2016).
40 Pseudochazara hippolyte (Esper, 1783) is a junior primary homonym of Papilio hyppolite Drury, 1782. The oldest available name for this taxon is Satyrus mercurius Staudinger, 1887.
41 Papilio afer Esper, 1783 is a junior primary homonym of Papilio afer Drury, 1782 (see Koçak 1981), as is Papilio afra Fabricius, 1787, because it differs only in gender. Therefore the oldest available name is Papilio phegea Borkhausen, 1788.
42 The name Papilio tithonus was first published in 1771 in Mantissa Plantarum Altera, not in 1767.
43 The name Hipparchia rossii was first published in November 1835, not in 1834.
44 The name Erebia dabanensis was published on 13 November 1872, not in 1871.
45 Recently, a population of Erebia was discovered in the Polar Urals and described as a new species, E. churkini Bogdanov, 2008, but is now considered a subspecies of Erebia callias (Tatarinov & Gorbunov, 2015). However, no further material is available, therefore it is tentatively considered as part of the European fauna. Erebia callias is a member of the tyndarus group (Albre et al. 2008) and ranges from the mountains of the Asian part of Russia and Mongolia to Colorado (USA).
46 The author of the name Papilio cassioides is Hohenwarth alone as indicated on page III of Reiner and Hohenwarth (1792), not Reiner and Hohenwarth.
47 Based on molecular data and differences in wing patterns Erebia cassioides has been split into three allopatric species (Schmitt et al. 2016). E. cassioides is limited to the eastern Alps, E. arvernensis is distributed in the western Alps, Cantabrian mountains and Pyrénées, while E. neleus is present in the mountains of the Balkan Peninsula and the southern Carpathians.
48 The name Erebia medusa polaris was first published in September 1861, not in 1871.
49 The year of publication of the name Erebia scipio by Boisduval is 1833, not 1832 (Cowan 1970).

Species richness of European butterfly families and subfamilies.

Family Subfamily Genera Species
Hesperiidae 13 47
Hesperiinae 6 11
Heteropterinae 2 3
Pyrginae 5 33
Lycaenidae 39 130
Aphnaeinae 1 1
Lycaeninae 1 13
Polyommatinae 30 98
Theclinae 7 18
Nymphalidae 41 246
Apaturinae 1 3
Charaxinae 1 1
Danainae 1 2
Heliconiinae 6 32
Libytheinae 1 1
Limenitidinae 2 5
Nymphalinae 8 37
Satyrinae 21 165
Papilionidae 5 15
Papilioninae 2 5
Parnassiinae 3 10
Pieridae 11 57
Coliadinae 3 18
Dismorphiinae 1 5
Pierinae 7 34
Riodinidae 1 1
Nemeobiinae 1 1
Total 21 110 496

Authors of currently valid European butterfly species (with a minimum of three described taxa).

Author Life data Nationality Species Period
Linnaeus, Carolus 1707–1778 Swedish 71 1758–1771
Poda von Neuhaus, Nicolaus (Nikolaus) 1723–1798 Austrian 4 1761
Scopoli, Giovanni Antonio 1723–1788 Italian 4 1763
Pallas, Peter Simon 1741–1811 German 8 1771
Schiffermüller, Johann Ignaz 1727–1806 Austrian 21 1775
Fabricius, Johan Christian 1745–1808 Danish 16 1775–1793
Rothenburg [alias Rottemburg], Siegmund Adrian von 1745–1797 German 8 1775
Esper, Eugen Johann Christoph 1742–1810 German 32 1777–1805
Bergsträsser, Johann Andreas Benignus 1732–1812 German 5 1779–1780
Knoch, August Wilhelm 1742–1818 German 3 1781–1783
Borkhausen, Moritz Balthasar 1760–1806 German 4 1788–1789
Hübner, Jacob 1761–1826 German 31 1790–1831
Thunberg, Carl Peter 1743–1828 Swedish 5 1791
Schneider, David Hinrich 1755–1826 German 3 1792–1794
Prunner, Leonhard von 17??–1830 German 8 1798
Hoffmansegg, Johann Centurius Graf von 1766–1849 German 6 1804–1806
Ochsenheimer, Ferdinand 1767–1822 German 4 1808–1816
Godart, Jean Baptiste 1775–1825 French 6 1819–1824
Freyer, Christian Friedrich 1794–1885 German 16 1828–1851
Boisduval, Jean Baptiste Alphonse Dechauffour de 1799–1879 French 13 1828–1848
Geyer, Carl 1802–1889 German 4 1828–1832
Klug, Johann Christoph Friedrich 1775–1856 German 4 1829–1834
Meigen, Johann Wilhelm 1764–1845 German 3 1829
Eversmann, Eduard Friedrich von 1794–1860 Russian 14 1832–1851
Rambur, Jules Pierre 1801–1870 French 10 1832–1839
Herrich-Schäffer, Gottlieb August Wilhelm 1799–1874 German 14 1844–1852
Zeller, Philipp Christoph 1808–1883 German 4 1847
Lederer, Julius 1821–1870 Austrian 3 1855–1864
Staudinger, Otto 1830–1900 German 17 1860–1901
Butler, Arthur Gardiner 1844–1925 British 6 1868–1898
Oberthür, Charles 1845–1924 French 9 1875–1910
Rebel, Hans 1861–1940 Austrian 5 1894–1916
Elwes, Henry John 1846–1922 British 3 1899–1900
Chapman, Thomas Algernon 1842–1921 British 3 1905–1920
Fruhstorfer, Hans 1866–1922 German 3 1908–1910
Verity, Ruggero 1883–1959 Italian 5 1921–1928
Kudrna, Otakar 1939– Czech 3 1976–1984
Brown, John 19??– British 3 1976–1981

Butterfly species excluded from the European list with explanations.

Turanana panagaea Distributed outside Europe in the Asian part of Turkey and replaced by Turanana taygetica in Europe (Hesselbarth et al. 1995; Coutsis 2005). [Junior subjective synonym of Lycaena endymion Gerhard, 1851; misspelled as Turanana panagea in Fauna Europaea]
(Herrich-Schäffer, 1851)
Polyommatus eleniae Considered conspecific with Polyommatus orphicus based on the equal haploid chromosome number and no differences in mitochondrial DNA – barcoding gene (Vishnevskaya et al. 2016).
Coutsis & De Prins, 2005
Polyommatus galloi According to the molecular study of Vila et al. (2010) P. galloi represents an isolated population of Polyommatus ripartii and is not considered as a separate species.
(Balletto & Toso, 1979)
Polyommatus menalcas Distributed outside Europe in Asian part of Turkey (Hesselbarth et al. 1995).
(Freyer, 1837)
Polyommatus pljushtchi Species status is based on erroneous sequences (opinion in Kudrna et al. (2011); Shapoval and Lukhtanov (2015).) Considered here as ssp. of Polyommatus damone (Eversmann, 1841).
Lukhtanov & Budashkin, 1993
Melitaea punica Distributed outside Europe in northern Africa (Toth et al. 2014).
Oberthür, 1876
Melitaea telona Distributed outside Europe in Levant (Toth et al. 2014).
Fruhstorfer, 1908
Pseudochazara mniszechii Distributed outside Europe in Asian part of Turkey (Hesselbarth et al. 1995). P. tisiphone, often considered as a subspecies of P. mniszechii, was shown not to be closely related to it (Verovnik and Wiemers 2016).
(Herrich-Schäffer, 1851)
Pseudochazara beroe Distributed outside Europe in Asian part of Turkey (Hesselbarth et al. 1995).
(Freyer, 1843)

List of the 14 European butterfly species that are affected by the gender agreement provision.

Name Original species epithet
Agriades pyrenaicus pyrenaica
Carcharodus tripolinus tripolina
Colias crocea croceus
Cupido decoloratus decolorata
Erebia aethiopella aethiopellus
Erebia albergana alberganus
Erebia montana montanus
Erebia stiria stirius
Hipparchia azorina azorinus
Hyponephele lupina lupinus
Kretania hesperica hespericus
Lycaena dimorpha dimorphus
Lycaena ottomana ottomanus
Tarucus balkanicus balkanica
Figure 4. 

Species richness of butterfly families in Europe.

Compared to the last version 2.6.2 of Fauna Europaea, nine species have been excluded from the list (Table 6). On the other hand, 15 species were added to the list. Another recently discovered species, Spialia rosae Hernández-Roldán, Dapporto, Dincă, Vicente & Vila, 2016, has already been added to the Fauna Europaea database.

Apart from the changes due to the gender agreement provision (Table 7), only three species names had to be changed due to new nomenclatural evidence: Pyrgus bellieri (Oberthür, 1910) to Pyrgus foulquieri (a name which had already been used in previous field guides), Proterebia afra (Fabricius, 1787) to Proterebia phegea (hopefully solving a longstanding controversy, see e.g., Jutzeler and Lafranchis 2011), and the mandatory change of Pseudochazara hippolyte (Esper, 1783) to Pseudochazara mercurius due to primary homonomy.

A larger number of changes concern the genus names. Most of them are in the family Lycaenidae, where 26 species changed their genus name, mainly based on the molecular study by Talavera et al. (2013), which substantially improved our knowledge of phylogenetic relationships of the subtribe Polyommatina. However, none of the genus names is new and many of them have already been used with the same species. In addition, four species formerly placed in the genus Argynnis were transferred into the genera Fabriciana and Speyeria, based on the study by De Moya et al. (2017). The former genus name had already been used previously for the same species, whereas the latter seems new to European lepidopterists, but is commonly used in North America. Although it could be argued that the change was avoidable by keeping a larger genus Argynnis, a solution originally also favoured by Simonsen et al. (2006), this would have meant to rename a large number of North American butterflies currently placed in the genus Speyeria, and was rejected by North American lepidopterists. Therefore, the recommended changes appear to cause the least changes on a global level and will hopefully contribute to a more consistent taxonomy of Holarctic Argynnini.

Finally, quite a number of minor changes have been implemented, which correct mistakes in names of authors, year of publication, or the incorrect use of parentheses for species that have changed generic combinations. An example is the change of year for 6 butterfly names due to a correction of the publication date of Linnaeus’ Fauna Svecica. Evenhuis (1997: 480) has shown convincingly that this edition was actually published on [14 November 1760], not “1761” as stated in the title page of the work and Bousquet (2016) also agrees with that year of publication.

Conclusions

Taking into account the many recent research findings, especially those with molecular methods, we think that the new taxonomy represents a step forward in stabilizing European butterfly taxonomy and nomenclature. Nevertheless, we have to note that some groups, e.g., the genera Euchloe, Callophrys, Pseudophilotes, Melitaea, and Hipparchia, as well as the subgenus Agrodiaetus of the genus Polyommatus are still in need of revision, which will certainly lead to additional changes in the future. Furthermore, we still have large knowledge gaps for species in other regions of the Palearctic region (especially in Central Asia), which might require changes in order to achieve a consistent taxonomy of Palearctic and Holarctic butterflies.

Acknowledgements

We thank Dirk Maes (Belgium) and his following collaborators for the country-level distribution data of European butterflies, which were assembled for another paper on national checklists and Red Lists (Maes et al. submitted): Jiří Beneš (CZ), Dimitri Brosens (BE), Stoyan Beshkov (BG), Simona Bonelli (IT), Jaroslaw Buszko (PL), Lisette Cantú Salazar (LU), Louis Francis Cassar (MT), Sue Collins (GB), Milan Djuric (RS), Goran Dusej (CH), Hallvard Elven (NO), Filip Franeta (RS), Patricia Garcia-Pereira (PT), Yurii Geryak (UA), Philippe Goffart (BE), Ádám Gór (HU), Ulrich Hiermann (AT), Helmut Höttinger (AT), Peter Huemer (AT), Predrag Jakšić (RS), Eddie John (CY), Henrik Kalivoda (SK), Vassiliki Kati (GR), Paul Kirkland (GB), Benjamin Komac (AD), Ádám Kőrösi (HU), Anatolij Kulak (BY), Mikko Kuussaari (FI), Lionel L’Hoste (LU), Suvad Lelo (BA), Xavier Mestdagh (LU), Nikola Micevski (MK), Iva Mihoci (HR), Sergiu Mihut (RO), Yeray Monasterio-León (ES), Dmitry V. Morgun (RU), Tomás Murray (IE), Per Stadel Nielsen (DK), Erling Ólafsson (IS), Erki Õunap (EE), Lazaros Pamperis (GR), Alois Pavlíčko (CZ), Lars B. Pettersson (SE), Serhiy Popov (UA), Miloš Popović (RS), Juha Pöyry (FI), Mike Prentice (GB), Nils Ryrholm (SE), Martina Šašić (HR), Nikolay Savenkov (LV), Josef Settele (DE), Marcin Sielezniew (PL), Sergey Sinev (RU), Constanti Stefanescu (ES), Giedrius Švitra (LT), Toomas Tammaru (EE), Anu Tiitsaar (EE), Elli Tzirkalli (CY), Olga Tzortzakaki (GR), Arne Lykke Viborg (DK), Martin S. Warren (GB), Irma Wynhoff (NL), and Konstantina Zografou (GR).

Our thanks also go to Ole Karsholt for his review which helped to improve the manuscript.

VL was supported by grant N 14-14-00541 from the Russian Science Foundation to the Zoological Institute of the Russian Academy of Sciences and ZF by grant 14-36098G from the Czech Science Foundation.

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Appendix 1

Evidence that the junior synonym Polyommatus ottomanus Lefèbvre, 1831 has been used to denote the taxon currently known as Lycaena ottomana (Lefèbvre, [1831]), in at least 25 works, published by at least 10 authors during the last 50 years and encompassing a span of not less than 10 years, and thus fulfilling the conditions of article 23.9.1.2 of the code in order to reverse the precedence of Lycaena legeri Freyer, 1830.

Already during the decades immediately following the publication of legeri Freyer, this name does not seem to have been used but as a subjective junior synonym of ottomanus Lefèbvre. The latter name was thought to represent the valid name and was first used in its original combination (Polyommatus ottomanus) and starting from the 20th century mostly in the combination of Chrysophanus ottomanus:

• Brullé (1832): Polyommatus ottomanus Lef.

• Herrich-Schäffer (1843): Polyomm. Ottomanus Lef.; synonym: Legeri

• Mann (1862): Polyommatus ottomanus Lef.

• Lang (1884): Polyommatus Ottomanus, Lefebrve [sic]; synonym: Legeri, Frr.

• Rebel (1903): Chrysophanus Ottomanus Lef.

• Spuler (1908): Chrysóphanus ottománus Lef.

• Courvoisier (1921): Chrysophanus ottomanus Lefebvre 1830; synonym: legeri Freyer, 1832

• Galvagni (1924): Chrysophanus ottomanus Lef.

• Rebel and Zerny (1934): Chrysophanus ottomanus Lef.

• Kanus (1963): Heodes (Chrysophanus) ottomanus Lef.

During the last 50 years we are not aware of any use of legeri Freyer, except as a subjective junior synonym of ottomanus Lefèbvre. The latter name was mostly used in the combination of Heodes ottomanus and later as Lycaena ottomanus or, due to the gender agreement rule of the code, as Lycaena ottomana:

1. Higgins and Riley (1970): Heodes ottomanus Lefèbvre, 1830

2. Higgins (1975): Heodes ottomanus Lefèbvre 1830

3. Higgins and Riley (1978): Heodes ottomanus Lefèbvre 1830

4. Schmidt-Koehl (1980): Heodes ottomanus Lefebvre, 1830

5. Krzywicki (1981): Heodes ottomanus Lefevre [sic]

6. Wiemers (1983): Heodes ottomanus ottomanus Lef.

7. Higgins and Riley (1983): Heodes ottomanus Lefèbvre, 1830

8. Kudrna (1986): Lycaena ottomanus Lefebvre, 1830

9. Jakšić (1988): Lycaena ottomanus Lefèbvre, 1830

10. Schaider and Jakšić (1989): Lycaena ottomanus Lef.

11. Hesselbarth et al. (1995): Lycaena ottomana (Lefebvre, [1830]); synonym: “Gen.IX. Lycaena. 182. Pap. Legeri” Freyer, C.F., [Dezember] 1830

12. Karsholt and Razowki (1996): Lycaena ottomanus (Lefèbvre, 1830)

13. Pamperis (1997): Heodes ottomanus

14. Jakšić (1998): Lycaena ottomanus Lefèbvre, 1830

15. Tolman and Lewington (1998): Lycaena ottomana (Lefèbvre, 1830)

16. Abadjiev (2001): Lycaena ottomana (Lefebvre, [1830])

17. Bozano and Weidenhoffer (2001): Lycaena ottomanus (Lefebvre, 1830); synonym: legeri Freyer, 1839

18. Mihoci et al. (2005): Lycaena ottomanus (Lefèbvre, 1830)

19. Coutsis and Ghavalas (2006): Lycaena ottomanus (Lefebvre, 1830)

20. Wagener (2006): Lycaena ottomanus (Lefebvre, 1830)

21. Settele et al. (2008): Lycaena ottomana (Lefebvre, 1830)

22. Tolman and Lewington (2008): Lycaena ottomana Lefèbvre, 1830

23. Pamperis (2009): Lycaena ottomanus

24. Van Swaay et al. (2010): Lycaena ottomana (Lefèbvre, 1830)

25. Tshikolovets (2011): Lycaena ottomana (Lefebvre, [1830]); synonym: legeri Freyer, 1839

26. Kemal and Koçak (2011): Lycaena (Heodes) ottomanus (Lefèbvre, [1830]); synonym: legeri Freyer, 1830

27. Kudrna et al. (2011): Lycaena ottomana (Lefebvre, 1831)

28. Koren et al. (2012): Lycaena ottomana (Lefèbvre, 1830)

29. Verovnik & Popović (2012): Lycaena ottomanus (Lefèbvre, 1830)

30. Kudrna et al. (2015): Lycaena ottomana (Lefebvre, 1831)

31. Çalişkan (2016): Lycaena ottomanus (Lefèbvre, [1830])

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