An updated checklist of the European Butterflies (Lepidoptera, Papilionoidea).

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 Polyommatusottomanus Lefèbvre, 1831, which is threatened by its senior synonym Lycaenalegeri Freyer, 1830, is declared a nomen protectum, thereby conserving its name in the current combination Lycaenaottomana.

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 (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 comprises 7 families (Table 1, Fig. 1) and includes the (skippers) and (moth butterflies). The skippers have previously been thought to represent the sister group to the butterflies and were often placed in a separate superfamily , but the molecular results indicate that the family is the sister to the remaining butterflies, which also include the small Neotropical family with only 36 species. Apart from the latter family, all butterfly families are represented on all continents except Antarctica, although most species of 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.

Table 1.

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

* global number of genera and species according to van Nieukerken et al. (2011)

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 . 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 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 ) was published by Seitz (1907–1909). At that time, already 90% of Europe’s butterfly species had PageBreakbeen 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 PageBreaktaxonomy 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 , 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 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 PageBreaklimited 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 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 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 )

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 PageBreakextinct, e.g., 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 , 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 (Verity, 1921), (Brown & Coutsis, 1978), (Stauder, 1921), Kudrna, 1984, Herrich-Schäffer, 1844, and 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 are as old as binominal nomenclature, because there is not even an agreement about the gender of the genus . 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 , which has turned out to represent another Asian species which is currently known as (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 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 ( to (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 Tutt, 1906 (type species: Ochsenheimer, 1808; currently known as (Ochsenheimer, 1808)), which appears to be a subjective synonym of the genus name Tutt, 1906 (type species: Lederer, 1858; currently known as (Lederer, 1858)). Both genus names were published in the same paper and PageBreakHemming (1967) was the first to note that should have precedence over , because Warren (1926) as the first reviser chose . However, the name 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 and needs to be substantially revised. However, molecular data are still missing for most of the (mainly Asian) species currently placed in , and therefore we suggest to postpone a rearrangement until better data become available.

Finally, one of us (GL) discovered that Lefèbvre was published in 1831 (and not in 1830) and therefore has to be regarded as a subjective junior synonym of Freyer, 1830. This would mean that the well-established name of the species currently known as (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., 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 Freyer as invalid and qualified as a nomen oblitum and declare the name 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.

Table 2.

Updated checklist of the butterflies of Europe.

Taxon	Original combination	Notes
Papilionidae
Papilioninae
Iphiclidespodalirius (Linnaeus, 1758)	Papilio podalirius
Iphiclidesfeisthamelii (Duponchel, 1832)	Papilio feisthamelii	1
Papilioalexanor Esper, 1800	Papilio alexanor
Papiliomachaon Linnaeus, 1758	Papilio machaon
Papiliohospiton Gené, 1839	Papilio hospiton	2
Parnassiinae
Parnassiusmnemosyne (Linnaeus, 1758)	Papilio mnemosyne
Parnassiusphoebus (Fabricius, 1793)	Papilio phoebus
Parnassiusapollo (Linnaeus, 1758)	Papilio apollo
Archonapollinus (Herbst, 1798)	Papilio apollinus
Zerynthiacerisy (Godart, [1824])	Thais cerisy
Zerynthiacretica (Rebel, 1904)	Thais cerisyi cretica
Zerynthiacaucasica (Lederer, 1864)	Thais cerisyi caucasica
Zerynthiarumina (Linnaeus, 1758)	Papilio rumina
Zerynthiapolyxena ([Denis & Schiffermüller], 1775)	Papilio polyxena
Zerynthiacassandra (Geyer, [1828])	Papilio cassandra	3
Hesperiidae
Heteropterinae
Heteropterusmorpheus (Pallas, 1771)	Papilio morpheus
Carterocephalussilvicola (Meigen, 1829)	Hesperia silvicola
Carterocephaluspalaemon (Pallas, 1771)	Papilio palaemon
Hesperiinae
Pelopidasthrax (Hübner, [1821])	Gegenes thrax
Borboborbonica (Boisduval, 1833)	Hesperia borbonica
Gegenespumilio (Hoffmansegg, 1804)	Papilio pumilio
Gegenesnostrodamus (Fabricius, 1793)	Hesperia nostrodamus
Ochlodessylvanus (Esper, 1777)	Papilio sylvanus
Hesperiacomma (Linnaeus, 1758)	Papilio comma
Thymelicuschristi Rebel, 1894	Thymelicus christi
Thymelicusacteon (Rottemburg, 1775)	Papilio acteon
Thymelicushyrax (Lederer, 1861)	Hesperia hyrax
Thymelicussylvestris (Poda, 1761)	Papilio sylvestris
Thymelicuslineola (Ochsenheimer, 1808)	Papilio lineola
Pyrginae
Spialiaphlomidis (Herrich-Schäffer, 1845)	Hesperia phlomidis
Spialiasertorius (Hoffmansegg, 1804)	Hesperia sertorius
Spialiatherapne (Rambur, 1832)	Hesperia therapne
Spialiarosae Hernández-Roldán, Dapporto, Dincă, Vicente & Vila, 2016	Spialia rosae	4
Spialiaorbifer (Hübner, [1823])	Papilio orbifer
Carcharodustripolinus (Verity, 1925)	Erynnis alceae tripolina	5
Carcharodusalceae (Esper, 1780)	Papilio alceae
Muschampiacribrellum (Eversmann, 1841)	Hesperia cribrellum
Muschampiatessellum (Hübner, [1803])	Papilio tessellum
Muschampiaproto (Ochsenheimer, 1808)	Papilio proto
Carcharoduslavatherae (Esper, 1783)	Papilio lavatherae
Carcharodusorientalis Reverdin, 1913	Carcharodus orientalis
Carcharodusfloccifera (Zeller, 1847)	Hesperia floccifera
Carcharodusstauderi Reverdin, 1913	Carcharodus stauderi
Carcharodusbaeticus (Rambur, 1839)	Spilothyrus baeticus
Erynnistages (Linnaeus, 1758)	Papilio tages
Erynnismarloyi (Boisduval, 1834)	Thanaos marloyi
Pyrgusmalvoides (Elwes & Edwards, 1897)	Hesperia malvoides
Pyrgusmalvae (Linnaeus, 1758)	Papilio malvae
Pyrguscarthami (Hübner, [1813])	Papilio carthami
Pyrgussidae (Esper, 1784)	Papilio sidae
Pyrguscentaureae (Rambur, 1839)	Hesperia centaureae
Pyrguscacaliae (Rambur, 1839)	Hesperia cacaliae
Pyrgusandromedae (Wallengren, 1853)	Syrichtus andromedae
Pyrgusserratulae (Rambur, 1839)	Hesperia serratulae
Pyrgusarmoricanus (Oberthür, 1910)	Syrichthus armoricanus
Pyrgusalveus (Hübner, [1803])	Papilio alveus
Pyrguswarrenensis (Verity, 1928)	Hesperia warrenensis
Pyrgusfoulquieri (Oberthür, 1910)	Syrichthus alveus foulquieri	6
Pyrgusonopordi (Rambur, 1839)	Hesperia onopordi
Pyrguscarlinae (Rambur, 1839)	Hesperia carlinae
Pyrguscirsii (Rambur, 1839)	Hesperia cirsii
Pyrguscinarae (Rambur, 1839)	Hesperia cinarae
Pieridae
Dismorphiinae
Leptideaduponcheli (Staudinger, 1871)	Leucophasia duponcheli
Leptideamorsei (Fenton, 1882)	Leptosia morsei
Leptideajuvernica Williams, 1946	Leptidea sinapis juvernica	7
Leptideasinapis (Linnaeus, 1758)	Papilio sinapis
Leptideareali Reissinger, 1990	Leptidea sinapis reali
Coliadinae
Gonepteryxrhamni (Linnaeus, 1758)	Papilio rhamni
Gonepteryxcleobule (Hübner, [1831])	Anteos cleobule	8
Gonepteryxcleopatra (Linnaeus, 1767)	Papilio cleopatra
Gonepteryxmaderensis C. Felder, 1862	Gonopteryx cleopatra maderensis
Gonepteryxfarinosa (Zeller, 1847)	Rhodocera farinosa
Catopsiliaflorella (Fabricius, 1775)	Papilio florella
Coliashyale (Linnaeus, 1758)	Papilio hyale
Coliasalfacariensis Ribbe, 1905	Colias hyale alfacariensis
Coliasphicomone (Esper, [1780])	Papilio phicomone
Coliasaurorina Herrich-Schäffer, 1850	Colias aurorina
Coliaschrysotheme (Esper, [1781])	Papilio chrysotheme
Coliaserate (Esper, [1805])	Papilio erate
Coliascrocea (Geoffroy, 1785)	Papilio croceus	5, 9
Coliasmyrmidone (Esper, [1781])	Papilio myrmidone
Coliascaucasica Staudinger, 1871	Colias myrmidone caucasica
Coliaspalaeno (Linnaeus, [1760])	Papilio palaeno	10
Coliastyche (Böber, 1812)	Papilio tyche
Coliashecla Lefèbvre, 1836	Colias hecla
Pierinae
Colotisevagore (Klug, 1829)	Pontia evagore
Aporiacrataegi (Linnaeus, 1758)	Papilio crataegi
Pontiachloridice (Hübner, [1813])	Papilio chloridice
Pontiacallidice (Hübner, [1800])	Papilio callidice
Pontiaedusa (Fabricius, 1777)	Papilio edusa
Pontiadaplidice (Linnaeus, 1758)	Papilio daplidice
Pieriskrueperi Staudinger, 1860	Pieris krueperi
Pierisbrassicae (Linnaeus, 1758)	Papilio brassicae
Pieriswollastoni (Butler, 1886)	Ganoris wollastoni
Pierischeiranthi (Hübner, [1808])	Papilio cheiranthi
Pierisrapae (Linnaeus, 1758)	Papilio rapae
Pierismannii (Mayer, 1851)	Pontia mannii
Pierisergane (Geyer, [1828])	Papilio ergane
Pierisbryoniae (Hübner, [1806])	Papilio bryoniae
Pierisnapi (Linnaeus, 1758)	Papilio napi
Pierisbalcana Lorković, [1969]	Pieris balcana	11
Euchloetagis (Hübner, [1804])	Papilio tagis
Euchloeeversi Stamm, 1963	Euchloe belemia eversi
Euchloegrancanariensis Acosta, 2008	Euchloe belemia grancanariensis
Euchloehesperidum Rothschild, 1913	Euchloe belemia hesperidum
Euchloebelemia (Esper, 1800)	Papilio belemia
Euchloeinsularis (Staudinger, 1861)	Anthocharis tagis insularis
Euchloecrameri Butler, 1869	Euchloe crameri
Euchloesimplonia (Freyer, 1829)	Pontia simplonia
Euchloeausonia (Hübner, [1804])	Papilio ausonia
Euchloecharlonia (Donzel, 1842)	Anthocharis charlonia
Euchloepenia (Freyer, 1851)	Pontia penia
Euchloebazae Fabiano, 1993	Euchloe charlonia bazae
Zegrispyrothoe (Eversmann, 1832)	Pontia pyrothoe
Zegriseupheme (Esper, [1804])	Papilio eupheme
Anthochariseuphenoides Staudinger, 1869	Anthocharis euphenoides
Anthochariscardamines (Linnaeus, 1758)	Papilio cardamines
Anthocharisgruneri Herrich-Schäffer, 1851	Anthocharis gruneri
Anthocharisdamone Boisduval, 1836	Anthocharis damone
Riodinidae
Nemeobiinae
Hamearislucina (Linnaeus, 1758)	Papilio lucina
Lycaenidae
Lycaeninae
Lycaenadimorpha (Staudinger, 1881)	Polyommatus dimorphus	5, 12
Lycaenahelle ([Denis & Schiffermüller], 1775)	Papilio helle
Lycaenaalciphron (Rottemburg, 1775)	Papilio alciphron
Lycaenathetis Klug, 1834	Lycaena thetis
Lycaenathersamon (Esper, 1784)	Papilio thersamon
Lycaenadispar ([Haworth], 1802)	Papilio dispar
Lycaenahippothoe (Linnaeus, [1760])	Papilio hippothoe	10
Lycaenacandens (Herrich-Schäffer, 1844)	Polyommatus candens
Lycaenaottomana (Lefèbvre, [1831])	Polyommatus ottomanus	5, 13
Lycaenableusei (Oberthür, 1884)	Polyommatus xanthe f. bleusei
Lycaenaphlaeas (Linnaeus, [1760])	Papilio phlaeas	10
Lycaenavirgaureae (Linnaeus, 1758)	Papilio virgaureae
Lycaenatityrus (Poda, 1761)	Papilio tityrus
Aphnaeinae
Cigaritisacamas (Klug, 1834)	Lycaena acamas	14
Theclinae
Theclabetulae (Linnaeus, 1758)	Papilio betulae
Favoniusquercus (Linnaeus, 1758)	Papilio quercus
Laeosopisroboris (Esper, [1793])	Papilio roboris	15
Tomaresballus (Fabricius, 1787)	Papilio ballus
Tomaresnogelii (Herrich-Schäffer, 1851)	Thecla nogelii
Tomarescallimachus (Eversmann, 1848)	Lycaena callimachus
Callophrysavis Chapman, 1909	Callophrys avis
Callophryssuaveola (Staudinger, 1881)	Thecla suaveola
Callophrysrubi (Linnaeus, 1758)	Papilio rubi
Callophryschalybeitincta Sovinsky, 1905	Callophrys rubi chalybeitincta
Neolycaenarhymnus (Eversmann, 1832)	Lycaena rhymnus
Satyriumpruni (Linnaeus, 1758)	Papilio pruni
Satyriumilicis (Esper, 1779)	Papilio ilicis
Satyriumesculi (Hübner, [1804])	Papilio esculi
Satyriumledereri (Boisduval, 1848)	Lycaena ledereri
Satyriumw-album (Knoch, 1782)	Papilio w-album
Satyriumspini ([Denis & Schiffermüller], 1775)	Papilio spini
Satyriumacaciae (Fabricius, 1787)	Papilio acaciae
Polyommatinae
Leptotespirithous (Linnaeus, 1767)	Papilio pirithous
Cyclyriuswebbianus (Brullé, 1839)	Polyommatus webbianus
Azanusubaldus (Stoll, 1782)	Papilio ubaldus
Azanusjesous (Guérin-Méneville, 1849)	Polyommatus jesous
Lampidesboeticus (Linnaeus, 1767)	Papilio boeticus
Cacyreusmarshalli Butler, 1898	Cacyreus marshalli
Celastrinaargiolus (Linnaeus, 1758)	Papilio argiolus
Tarucustheophrastus (Fabricius, 1793)	Hesperia theophrastus
Tarucusbalkanicus (Freyer, 1844)	Lycaena balkanica	5
Phengarisalcon ([Denis & Schiffermüller], 1775)	Papilio alcon
Phengarisarion (Linnaeus, 1758)	Papilio arion
Phengaristeleius (Bergsträsser, 1779)	Papilio teleius
Phengarisnausithous (Bergsträsser, 1779)	Papilio nausithous
Turananataygetica (Rebel, 1902)	Lycaena panagaea taygetica
Pseudophilotesbavius (Eversmann, 1832)	Lycaena bavius
Pseudophilotesbarbagiae De Prins & van der Poorten, 1982	Pseudophilotes barbagiae
Pseudophilotesabencerragus (Pierret, 1837)	Argus abencerragus
Pseudophilotespanoptes (Hübner, [1813])	Papilio panoptes
Pseudophilotesvicrama (Moore, 1865)	Polyommatus vicrama
Pseudophilotesbaton (Bergsträsser, 1779)	Papilio baton
Scolitantidesorion (Pallas, 1771)	Papilio orion
Praephilotesanthracias (Christoph, 1877)	Lycaena anthracias
Iolanaiolas (Ochsenheimer, 1816)	Lycaena iolas
Iolanadebilitata (Schultz, 1905)	Lycaena jolas var. debilitata	16
Glaucopsychemelanops (Boisduval, 1828)	Polyommatus melanops
Glaucopsychepaphos Chapman, 1920	Glaucopsyche paphos
Glaucopsychealexis (Poda, 1761)	Papilio alexis
Zizeeriaknysna (Trimen, 1862)	Lycaena knysna
Zizeeriakarsandra (Moore, 1865)	Polyommatus karsandra
Tongeiafischeri (Eversmann, 1843)	Lycaena fischeri
Cupidoargiades (Pallas, 1771)	Papilio argiades
Cupidodecoloratus (Staudinger, 1886)	Lycaena argiades decolorata	5
Cupidoalcetas (Hoffmansegg, 1804)	Papilio alcetas
Cupidoosiris (Meigen, 1829)	Polyommatus osiris
Cupidominimus (Fuessly, 1775)	Papilio minimus
Cupidolorquinii (Herrich-Schäffer, 1850)	Lycaena lorquinii	17
Luthrodesgalba (Lederer, 1855)	Lycaena galba	18
Freyeriatrochylus (Freyer, 1844)	Lycaena trochylus	18,19
Plebejusargus (Linnaeus, 1758)	Papilio argus
Plebejusidas (Linnaeus, [1760])	Papilio idas	10
Plebejusbellieri (Oberthür, 1910)	Lycaena bellieri
Plebejusargyrognomon (Bergsträsser, 1779)	Papilio argyrognomon
Agriadesorbitulus (Prunner, 1798)	Papilio orbitulus	18
Agriadesoptilete (Knoch, 1781)	Papilio optilete	18
Agriadespyrenaicus (Boisduval, 1840)	Lycaena orbitulus var. pyrenaica	5, 18
Agriadesdardanus (Freyer, 1843)	Lycaena dardanus	18
Agriadeszullichi Hemming, 1933	Agriades zullichi	18
Agriadesglandon (Prunner, 1798)	Papilio glandon	18
Agriadesaquilo (Boisduval, 1832)	Argus aquilo	18
Plebejidealoewii (Zeller, 1847)	Lycaena loewii	18
Eumedoniaeumedon (Esper, 1780)	Papilio eumedon	18
Kretaniapsylorita (Freyer, 1845)	Lycaena psylorita	18
Kretaniahesperica (Rambur, 1839)	Polyommatus hespericus	5, 18
Kretaniaeurypilus (Freyer, 1851)	Lycaena eurypilus	18
Kretaniatrappi (Verity, 1927)	Lycaena trappi	18
Kretaniasephirus (Frivaldszky, 1835)	Lycaena sephirus	18
Kretaniapylaon (Fischer, 1832)	Lycaena pylaon	18
Cyanirissemiargus (Rottemburg, 1775)	Papilio semiargus
Glabroculuscyane (Eversmann, 1837)	Lycaena cyane	18
Ariciamorronensis (Ribbe, 1910)	Lycaena idas morronensis
Ariciaanteros (Freyer, 1838)	Lycaena anteros
Ariciacramera (Eschscholtz, 1821)	Lycaena cramera
Aricianicias (Meigen, 1829)	Polyommatus nicias	20
Ariciaartaxerxes (Fabricius, 1793)	Hesperia artaxerxes
Ariciamontensis Verity, 1928	Aricia medon montensis
Ariciaagestis ([Denis & Schiffermüller], 1775)	Papilio agestis
Neolysandracoelestina (Eversmann, 1843)	Lycaena coelestina	18
Lysandrahispana (Herrich-Schäffer, 1851)	Lycaena coridon var. hispana	18
Lysandracorydonius (Herrich-Schäffer, 1852)	Lycaena coridon corydonius	18
Lysandrabellargus (Rottemburg, 1775)	Papilio bellargus	18
Lysandracoridon (Poda, 1761)	Papilio coridon	18
Lysandracaelestissima (Verity, 1921)	Agriades coridon caelestissima	18
Lysandraalbicans (Gerhard, 1851)	Lycaena coridon var. albicans	18
Polyommatusescheri (Hübner, [1823])	Papilio escheri
Polyommatusthersites (Cantener, 1835)	Argus thersites
Polyommatusdaphnis ([Denis & Schiffermüller], 1775)	Papilio daphnis
Polyommatusamandus (Schneider, 1792)	Papilio amandus
Polyommatusgolgus (Hübner, [1813])	Papilio golgus
Polyommatusnivescens (Keferstein, 1851)	Lycaena dorylas var. nivescens
Polyommatusdorylas ([Denis & Schiffermüller], 1775)	Papilio dorylas
Polyommatuscelina (Austaut, 1879)	Lycaena celina	21
Polyommatusicarus (Rottemburg, 1775)	Papilio icarus
Polyommatuseros (Ochsenheimer, 1808)	Papilio eros
Polyommatusdamon ([Denis & Schiffermüller], 1775)	Papilio damon
Polyommatusdamone (Eversmann, 1841)	Lycaena damone
Polyommatusdamocles (Herrich-Schäffer, 1844)	Lycaena damocles
Polyommatusadmetus (Esper, 1783)	Papilio admetus
Polyommatusripartii (Freyer, 1830)	Lycaena ripartii
Polyommatusnephohiptamenos (Brown & Coutsis, 1978)	Agrodiaetus nephohiptamenos
Polyommatusiphigenia (Herrich-Schäffer, 1847)	Lycaena iphigenia
Polyommatusvioletae (Gómez-Bustillo, Expósito & Martínez, 1979)	Agrodiaetus violetae
Polyommatusfulgens (Sagarra, 1925)	Hirsutina dolus r. fulgens	22
Polyommatusfabressei (Oberthür, 1910)	Lycaena rippertii r. fabressei
Polyommatusdolus (Hübner, [1823])	Papilio dolus
Polyommatushumedasae (Toso & Balletto, 1976)	Agrodiaetus humedasae
Polyommatustimfristos Lukhtanov, Vishnevskaya & Shapoval, 2016	Polyommatus timfristos	23
Polyommatusorphicus Kolev, 2005	Polyommatus orphicus
Polyommatusaroaniensis (Brown, 1976)	Agrodiaetus alcestis aroaniensis
Nymphalidae
Limenitidinae
Neptissappho (Pallas, 1771)	Papilio sappho
Neptisrivularis (Scopoli, 1763)	Papilio rivularis
Limenitisreducta Staudinger, 1901	Limenitis camilla reducta
Limenitispopuli (Linnaeus, 1758)	Papilio populi
Limenitiscamilla (Linnaeus, 1764)	Papilio camilla
Heliconiinae
Issorialathonia (Linnaeus, 1758)	Papilio lathonia
Issoriaeugenia (Eversmann, 1847)	Argynnis eugenia
Brenthishecate ([Denis & Schiffermüller], 1775)	Papilio hecate
Brenthisino (Rottemburg, 1775)	Papilio ino
Brenthisdaphne ([Denis & Schiffermüller], 1775)	Papilio daphne
Argynnispaphia (Linnaeus, 1758)	Papilio paphia
Argynnispandora ([Denis & Schiffermüller], 1775)	Papilio pandora
Argynnislaodice (Pallas, 1771)	Papilio laodice
Speyeriaaglaja (Linnaeus, 1758)	Papilio aglaja	24
Fabricianaelisa (Godart, 1823)	Argynnis elisa	24
Fabriciananiobe (Linnaeus, 1758)	Papilio niobe	24
Fabricianaadippe ([Denis & Schiffermüller], 1775)	Papilio adippe	24
Boloriaeunomia (Esper, 1800)	Papilio eunomia	25
Boloriagraeca (Staudinger, 1870)	Argynnis pales graeca
Boloriapales ([Denis & Schiffermüller], 1775)	Papilio pales
Boloriaalaskensis (Holland, 1900)	Argynnis alaskensis
Bolorianapaea (Hoffmansegg, 1804)	Papilio napaea
Boloriaaquilonaris (Stichel, 1908)	Argynnis aquilonaris
Boloriatritonia (Böber, 1812)	Papilio tritonia
Boloriapolaris (Boisduval, 1828)	Argynnis polaris
Boloriathore (Hübner, [1804])	Papilio thore	26
Boloriaselene ([Denis & Schiffermüller], 1775)	Papilio selene
Boloriaeuphrosyne (Linnaeus, 1758)	Papilio euphrosyne
Boloriadia (Linnaeus, 1767)	Papilio dia
Boloriaimproba (Butler, 1877)	Argynnis improba
Boloriafrigga (Thunberg, 1791)	Papilio frigga	27
Boloriafreija (Thunberg, 1791)	Papilio freija	27
Boloriaselenis (Eversmann, 1837)	Argynnis selenis
Boloriaoscarus (Eversmann, 1844)	Argynnis oscarus
Boloriatitania (Esper, [1793])	Papilio titania
Boloriachariclea (Schneider, 1794)	Papilio chariclea
Boloriaangarensis (Erschoff, 1870)	Argynnis angarensis
Apaturinae
Apaturairis (Linnaeus, 1758)	Papilio iris
Apaturametis Freyer, 1829	Apatura metis
Apaturailia ([Denis & Schiffermüller], 1775)	Papilio ilia
Nymphalinae
Araschnialevana (Linnaeus, 1758)	Papilio levana
Vanessavirginiensis (Drury, 1773)	Papilio cardui virginiensis
Vanessacardui (Linnaeus, 1758)	Papilio cardui
Vanessavulcania Godart, 1819	Vanessa vulcania
Vanessaatalanta (Linnaeus, 1758)	Papilio atalanta
Aglaisio (Linnaeus, 1758)	Papilio io
Aglaisurticae (Linnaeus, 1758)	Papilio urticae
Aglaisichnusa (Hübner, [1824])	Papilio ichnusa	28
Polygoniaegea (Cramer, 1775)	Papilio egea
Polygoniac-album (Linnaeus, 1758)	Papilio c-album
Nymphalisvaualbum ([Denis & Schiffermüller], 1775)	Papilio vau album
Nymphalispolychloros (Linnaeus, 1758)	Papilio polychloros
Nymphalisxanthomelas ([Denis & Schiffermüller], 1775)	Papilio xanthomelas
Nymphalisantiopa (Linnaeus, 1758)	Papilio antiopa
Hypolimnasmisippus (Linnaeus, 1764)	Papilio misippus
Euphydryasdesfontainii (Godart, 1819)	Papilio desfontainii
Euphydryasaurinia (Rottemburg, 1775)	Papilio aurinia
Euphydryascynthia ([Denis & Schiffermüller], 1775)	Papilio cynthia
Euphydryasiduna (Dalman, 1816)	Melitaea iduna
Euphydryasmaturna (Linnaeus, 1758)	Papilio maturna
Euphydryasintermedia (Ménétriés, 1859)	Melitaea maturna intermedia
Melitaeatrivia ([Denis & Schiffermüller], 1775)	Papilio trivia
Melitaeadidyma (Esper, 1778)	Papilio didyma
Melitaeaarduinna (Esper, 1783)	Papilio arduinna
Melitaeaaetherie (Hübner, [1826])	Papilio aetherie
Melitaeaphoebe ([Denis & Schiffermüller], 1775)	Papilio phoebe
Melitaeaornata Christoph, 1893	Melitaea phoebe ornata	29
Melitaeacinxia (Linnaeus, 1758)	Papilio cinxia
Melitaeadiamina (Lang, 1789)	Papilio diamina
Melitaeaceladussa Fruhstorfer, 1910	Melitaea athalia celadussa	30
Melitaeadeione (Geyer, [1832])	Papilio deione
Melitaeabritomartis Assmann, 1847	Melitaea britomartis
Melitaeaathalia (Rottemburg, 1775)	Papilio athalia
Melitaeavaria Herrich-Schäffer, 1851	Melitaea varia	31
Melitaeaparthenoides Keferstein, 1851	Melitaea athalia parthenoides
Melitaeaaurelia Nickerl, 1850	Melitaea aurelia
Melitaeaasteria Freyer, 1828	Melitaea asteria
Libytheinae
Libytheaceltis (Laicharting, 1782)	Papilio celtis
Danainae
Danausplexippus (Linnaeus, 1758)	Papilio plexippus
Danauschrysippus (Linnaeus, 1758)	Papilio chrysippus
Charaxinae
Charaxesjasius (Linnaeus, 1767)	Papilio jasius
Satyrinae
Coenonymphaphryne (Pallas, 1771)	Papilio phryne
Coenonymphaoedippus (Fabricius, 1787)	Papilio oedippus
Coenonymphadorus (Esper, 1782)	Papilio dorus
Coenonymphathyrsis (Freyer, 1845)	Hipparchia thyrsis
Coenonymphapamphilus (Linnaeus, 1758)	Papilio pamphilus
Coenonymphatullia (Müller, 1764)	Papilio tullia
Coenonympharhodopensis Elwes, 1900	Coenonympha tiphon rhodopensis
Coenonymphaamaryllis (Stoll, 1782)	Papilio amaryllis
Coenonymphaglycerion (Borkhausen, 1788)	Papilio glycerion
Coenonymphacorinna (Hübner, [1804])	Papilio corinna
Coenonymphaleander (Esper, 1784)	Papilio leander
Coenonymphahero (Linnaeus, [1760])	Papilio hero	10
Coenonymphagardetta (Prunner, 1798)	Papilio gardetta
Coenonymphaorientalis Rebel, 1909	Coenonympha arcania var. orientalis	32
Coenonymphaarcania (Linnaeus, [1760])	Papilio arcania	10
Kiriniaroxelana (Cramer, 1777)	Papilio roxelana
Kiriniaclimene (Esper, 1783)	Papilio climene
Lopingaachine (Scopoli, 1763)	Papilio achine
Parargexiphia (Fabricius, 1775)	Papilio xiphia
Parargexiphioides Staudinger, 1871	Pararge xiphia xiphioides
Parargeaegeria (Linnaeus, 1758)	Papilio aegeria
Lasiommatamaera (Linnaeus, 1758)	Papilio maera
Lasiommatadeidamia (Eversmann, 1851)	Hipparchia deidamia
Lasiommatapetropolitana (Fabricius, 1787)	Papilio maera petropolitana
Lasiommataparamegaera (Hübner, [1824])	Papilio paramegaera
Lasiommatamegera (Linnaeus, 1767)	Papilio megera
Melanargiarussiae (Esper, 1783)	Papilio russiae
Melanargialarissa (Geyer, [1828])	Papilio larissa
Melanargialachesis (Hübner, 1790)	Papilio lachesis
Melanargiagalathea (Linnaeus, 1758)	Papilio galathea
Melanargiaines (Hoffmansegg, 1804)	Papilio ines
Melanargiaarge (Sulzer, 1776)	Papilio arge
Melanargiapherusa (Boisduval, 1833)	Arge pherusa
Melanargiaoccitanica (Esper, [1793])	Papilio arge occitanica
Hipparchiafatua Freyer, 1843	Hipparchia fatua	33
Hipparchiastatilinus (Hufnagel, 1766)	Papilio statilinus
Hipparchiatilosi Manil, 1984	Hipparchia wyssii tilosi
Hipparchiabacchus (Higgins, 1967)	Pseudotergumia wyssii bacchus
Hipparchiawyssii (Christ, 1889)	Satyrus fidia wyssii
Hipparchiatamadabae Owen & Smith, 1992	Hipparchia wyssi tamadabae
Hipparchiagomera (Higgins, 1967)	Pseudotergumia wyssii gomera
Hipparchiafidia (Linnaeus, 1767)	Papilio fidia
Hipparchianeomiris (Godart, 1823)	Satyrus neomiris	34
Hipparchiaautonoe (Esper, 1783)	Papilio autonoe
Hipparchiahermione (Linnaeus, 1764)	Papilio hermione
Hipparchiasyriaca (Staudinger, 1871)	Satyrus hermione syriaca
Hipparchiafagi (Scopoli, 1763)	Papilio fagi
Hipparchiamersina (Staudinger, 1871)	Satyrus semele mersina
Hipparchiamiguelensis (Le Cerf, 1935)	Satyrus azorinus miguelensis
Hipparchiaazorina (Strecker, 1899)	Satyrus azorinus	5, 35
Hipparchiasenthes (Fruhstorfer, 1908)	Eumenis semele senthes
Hipparchiamaderensis (Bethune-Baker, 1891)	Satyrus semele maderensis
Hipparchiasemele (Linnaeus, 1758)	Papilio semele
Hipparchiablachieri (Fruhstorfer, 1908)	Eumenis semele blachieri
Hipparchiaaristaeus (Bonelli, 1826)	Papilio aristaeus
Hipparchiavolgensis (Mazokhin-Porshnyakov, 1952)	Satyrus semele volgensis
Hipparchianeapolitana (Stauder, 1921)	Satyrus neapolitana
Hipparchialeighebi Kudrna, 1976	Hipparchia semele leighebi
Hipparchiapellucida (Stauder, 1924)	Satyrus semele pellucida	36
Hipparchiasbordonii Kudrna, 1984	Hipparchia sbordonii
Hipparchiacypriensis (Holik, 1949)	Satyrus semele cypriensis
Hipparchiacretica (Rebel, 1916)	Satyrus semele cretica
Hipparchiachristenseni Kudrna, 1977	Hipparchia christenseni
Minoisdryas (Scopoli, 1763)	Papilio dryas
Brintesiacirce (Fabricius, 1775)	Papilio circe
Arethusanaarethusa ([Denis & Schiffermüller], 1775)	Papilio arethusa
Oeneistarpeia (Pallas, 1771)	Papilio tarpeia
Oeneisbore (Schneider, 1792)	Papilio bore
Oeneisammon Elwes, 1899	Oeneis bore var. ammon	37
Oeneismelissa (Fabricius, 1775)	Papilio melissa
Oeneismagna Graeser, 1888	Oeneis jutta magna
Oeneisjutta (Hübner, [1806])	Papilio jutta
Oeneisnorna (Thunberg, 1791)	Papilio norna
Oeneispolixenes (Fabricius, 1775)	Papilio polixenes
Oeneisglacialis (Moll, 1785)	Papilio glacialis	38
Satyrusferula (Fabricius, 1793)	Papilio ferula
Satyrusvirbius Herrich-Schäffer, 1844	Satyrus virbius
Satyrusactaea (Esper, 1781)	Papilio actaea
Chazarabriseis (Linnaeus, 1764)	Papilio briseis
Chazaraprieuri (Pierret, 1837)	Satyrus prieuri
Chazarapersephone (Hübner, [1805])	Papilio persephone
Pseudochazarageyeri (Herrich-Schäffer, 1846)	Satyrus geyeri
Pseudochazaragraeca (Staudinger, 1870)	Satyrus pelopea graeca
Pseudochazaraamymone Brown, 1976	Pseudochazara amymone
Pseudochazaraanthelea (Hübner, [1824])	Papilio anthelea
Pseudochazaraamalthea (Frivaldszky, 1845)	Hipparchia amalthea	39
Pseudochazarawilliamsi (Romei, 1927)	Satyrus hippolyte williamsi
Pseudochazaraeuxina (Kuznetsov, 1909)	Hipparchia euxina
Pseudochazaramercurius (Staudinger, 1887)	Satyrus mercurius	40
Pseudochazaracingovskii (Gross, 1973)	Satyrus sintenisi cingovskii
Pseudochazaratisiphone Brown, [1981]	Pseudochazara cingovskii tisiphone	39
Pseudochazaraorestes De Prins & van der Poorten, 1981	Pseudochazara orestes
Ypthimaasterope (Klug, 1832)	Hipparchia asterope
Proterebiaphegea (Borkhausen, 1788)	Papilio phegea	41
Hyponephelehuebneri Koçak, 1980	Hyponephele huebneri
Hyponephelelycaon (Kühn, 1774)	Papilio lycaon
Hyponephelelupina (Costa, 1836)	Satyrus lupinus	5
Aphantopushyperantus (Linnaeus, 1758)	Papilio hyperantus
Pyroniacecilia (Vallantin, 1894)	Epinephele ida cecilia
Pyroniatithonus (Linnaeus, 1771)	Papilio tithonus	42
Pyroniabathseba (Fabricius, 1793)	Papilio bathseba
Maniolajurtina (Linnaeus, 1758)	Papilio jurtina
Maniolanurag (Ghiliani, 1852)	Satyrus nurag
Maniolachia Thomson, 1987	Maniola chia
Maniolamegala (Oberthür, 1909)	Epinephele janira megala
Maniolacypricola (Graves, 1928)	Epinephele cypricola
Maniolatelmessia (Zeller, 1847)	Hipparchia telmessia
Maniolahalicarnassus Thomson, 1990	Maniola halicarnassus
Erebiaedda Ménétriés, 1851	Erebia edda
Erebiafasciata Butler, 1868	Erebia fasciata
Erebiadiscoidalis (Kirby, 1837)	Hipparchia discoidalis
Erebiarossii (Curtis, 1835)	Hipparchia rossii	43
Erebiacyclopius (Eversmann, 1844)	Hipparchia cyclopius
Erebiaembla (Thunberg, 1791)	Papilio embla
Erebiadisa (Thunberg, 1791)	Papilio disa
Erebiameolans (Prunner, 1798)	Papilio meolans
Erebiadabanensis Erschoff, 1872	Erebia dabanensis	44
Erebiajeniseiensis Trybom, 1877	Erebia ligea jeniseiensis
Erebiaclaudina (Borkhausen, 1789)	Papilio claudina
Erebiamanto ([Denis & Schiffermüller], 1775)	Papilio manto
Erebiaottomana Herrich-Schäffer, 1847	Erebia dromus ottomana
Erebiahispania Butler, 1868	Erebia hispania
Erebiarondoui Oberthür, 1908	Erebia rondoui
Erebiacallias Edwards, 1871	Erebia callias	45
Erebiatyndarus (Esper, 1781)	Papilio tyndarus
Erebiacassioides (Hohenwarth, 1792)	Papilio cassioides	46
Erebianivalis Lorković & Lesse, 1954	Erebia nivalis
Erebianeleus (Freyer, 1832)	Hipparchia neleus	47
Erebiacalcarius Lorković, 1953	Erebia tyndarus calcarius
Erebiaarvernensis Oberthür, 1908	Erebia tyndarus arvernensis	47
Erebiaoeme (Hübner, [1804])	Papilio oeme
Erebiagorge (Hübner, [1804])	Papilio gorge
Erebiasthennyo Graslin, 1850	Erebia sthennyo
Erebiapandrose (Borkhausen, 1788)	Papilio pandrose
Erebiaeriphyle (Freyer, 1836)	Hipparchia eriphyle
Erebiaepistygne (Hübner, [1819])	Papilio epistygne
Erebiaeuryale (Esper, 1805)	Papilio euryale
Erebiapalarica Chapman, 1905	Erebia palarica
Erebialigea (Linnaeus, 1758)	Papilio ligea
Erebiapluto (Prunner, 1798)	Papilio pluto
Erebiaaethiopellus (Hoffmansegg, 1806)	Papilio aethiopellus
Erebiagorgone Boisduval, 1833	Erebia gorgone
Erebiarhodopensis Nicholl, 1900	Erebia gorgone rhodopensis
Erebiamnestra (Hübner, [1804])	Papilio mnestra
Erebiaalbergana (Prunner, 1798)	Papilio alberganus	5
Erebiasudetica Staudinger, 1861	Erebia melampus sudetica
Erebiamelampus (Fuessly, 1775)	Papilio melampus
Erebiatriarius (Prunner, 1798)	Papilio triarius
Erebiapolaris Staudinger, 1861	Erebia medusa var. polaris	48
Erebiamedusa ([Denis & Schiffermüller], 1775)	Papilio medusa
Erebiaaethiops (Esper, 1777)	Papilio aethiops
Erebiapharte (Hübner, [1804])	Papilio pharte
Erebiachristi Rätzer, 1890	Erebia christi
Erebiaorientalis Elwes, 1900	Erebia epiphron orientalis
Erebiaepiphron (Knoch, 1783)	Papilio epiphron
Erebiaflavofasciata Heyne, 1895	Erebia flavofasciata
Erebiamontana (Prunner, 1798)	Papilio montanus	5
Erebiastyx (Freyer, 1834)	Hipparchia styx
Erebiastiria (Godart, [1824])	Satyrus stirius	5
Erebiascipio Boisduval, 1833	Erebia scipio	49
Erebiapronoe (Esper, 1780)	Papilio pronoe
Erebiamelas (Herbst, 1796)	Papilio melas
Erebialefebvrei (Boisduval, 1828)	Satyrus lefebvrei
Erebiazapateri Oberthür, 1875	Erebia zapateri
Erebianeoridas (Boisduval, 1828)	Satyrus neoridas

Table 4.

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

Figure 4.

Species richness of butterfly families in Europe.

Table 5.

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

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, Hernández-Roldán, Dapporto, Dincă, Vicente & Vila, 2016, has already been added to the Fauna Europaea database.

Table 6.

Butterfly species excluded from the European list with explanations.

Turanana panagaea	Distributed outside Europe in the Asian part of Turkey and replaced by Turananataygetica in Europe (Hesselbarth et al. 1995; Coutsis 2005). [Junior subjective synonym of Lycaenaendymion Gerhard, 1851; misspelled as Turananapanagea in Fauna Europaea]
(Herrich-Schäffer, 1851)
Polyommatus eleniae	Considered conspecific with Polyommatusorphicus 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 Polyommatusripartii 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 Polyommatusdamone (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)

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: (Oberthür, 1910) to (a name which had already been used in previous field guides), (Fabricius, 1787) to (hopefully solving a longstanding controversy, see e.g., Jutzeler and Lafranchis 2011), and the mandatory change of (Esper, 1783) to due to primary homonomy.

Table 7.

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

A larger number of changes concern the genus names. Most of them are in the family , 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 . 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 were transferred into the genera and , 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 , 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 , 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 .

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 , , , , and , as well as the subgenus of the genus 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.

Table 3.

Annotations to the updated checklist of the butterflies of Europe.

1	Iphiclidesfeisthamelii 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 Zerynthiacassandra 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	Spialiarosae 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 Syrichtusalveus f. foulquieri and Syrichtusalveus f. bellieri were published simultaneously (Oberthür, 1910), the name used by the first reviser (i. e. Rebel 1914), Pyrgusfoulquieri, should be used in accordance with Art. 24.2.1 and Art. 24.2.2 (ICZN 1999).
7	Recent studies have shown that Leptideareali 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 Anteoscleobule is 1831, not 1830 (the original plate [79], published in 1824, carried no names).
9	The name Papiliocroceus 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 Pierisbalcana 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 Lycaenadimorphairghiza 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 Polyommatusottomanus 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 Papilioroboris was first published in 1793, not 1789 (Lamas 2013).
16	Iolanadebilitata 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 Cupidolorquinii 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 Lycaenatrochylus is 1844, not 1845 (Tremewan 1988, Olivier 2000).
20	The year of the publication of the name Polyommatusnicias is ante September 1829, not 1830 (Griffin 1931).
21	Polyommatuscelina 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	Polyommatustimfristos 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 Papilioeunomia was first published in 1800, not 1799 (Poche, 1938).
26	The name Papiliothore was first published in 1804, not 1803 (Hemming 1937).
27	Description of Boloriafreija and Boloriafrigga must be credited to Thunberg, not to Becklin (Thunberg wrote Becklin's dissertation), see Karsholt and Nielsen (1986).
28	Papilioichnusa was first described by Hübner (ante 23 December) 1824. Vanessaichnusa 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 Melitaeaphoebe 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	Melitaeaceladussa 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 Melitaeaparthenievar.nevadensis Spuler, 1901, currently regarded as a junior subjective synonym of Melitaeaparthenoides Keferstein, 1851.
31	Melitaeavaria was first described by Herrich-Schäffer (1851) in Systematische Bearbeitung der Schmetterlinge von Europa Vol. 6(48): 2 (Hemming 1937). Melitaeaparthenievar.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 Hipparchiafatua was first published in 1843, not 1844 (Olivier 2000).
34	The name Satyrusneomiris was first published in 1823, not 1822. Satyrusneomiris 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 Satyrusazorinus was first published in 1899, not 1898.
36	The name Satyrussemelepellucida was first published on 15 May 1924, not in 1923.
37	Oeneisammon is present in Europe in the Polar Urals (Tsvetkov 2006).
38	The name Papilioglacialis was first published in 1785, not 1783.
39	Based on differentiation in mtDNA (barcodes) and differences in morphology, Pseudochazaraamalthea and P.tisiphone are considered separate species from allopatric P.anthelea and P.mniszechii respectively (Verovnik and Wiemers 2016).
40	Pseudochazarahippolyte (Esper, 1783) is a junior primary homonym of Papiliohyppolite Drury, 1782. The oldest available name for this taxon is Satyrusmercurius Staudinger, 1887.
41	Papilioafer Esper, 1783 is a junior primary homonym of Papilioafer Drury, 1782 (see Koçak 1981), as is Papilioafra Fabricius, 1787, because it differs only in gender. Therefore the oldest available name is Papiliophegea Borkhausen, 1788.
42	The name Papiliotithonus was first published in 1771 in Mantissa Plantarum Altera, not in 1767.
43	The name Hipparchiarossii was first published in November 1835, not in 1834.
44	The name Erebiadabanensis 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 Erebiacallias (Tatarinov & Gorbunov, 2015). However, no further material is available, therefore it is tentatively considered as part of the European fauna. Erebiacallias 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 Papiliocassioides 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 Erebiacassioides 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 Erebiamedusapolaris was first published in September 1861, not in 1871.
49	The year of publication of the name Erebiascipio by Boisduval is 1833, not 1832 (Cowan 1970).