Toward a revised checklist of the Western Palearctic butterflies, hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea)
Part VI: Rationale and framework for the Lycaenidae (addendum part II)
Submitted: 12.ix.2026 | Accepted: 15.ix.2026 | Published online: 30.ix.2026.
DOI: 10.5281/zenodo.22302254

Michel Taymans¹ | Sylvain Cuvelier²
0
¹ Clos du Moulin Royal 2 bte 02, B-6900 Marche-en-Famenne, Belgium. michel.taymans@hotmail.com
² Diamantstraat 4, B-8900 Ieper, Belgium. sylvain.cuvelier@telenet.be

Abstract
This second review of taxonomic and nomenclatural issues in Lycaenidae examines the status of family-group names in the light of the International Code of Zoological Nomenclature (ICZN), historical classification schemes, recent phylogenetic evidence, and current usage. Emphasis is placed on the classification of subfamilies and tribes, for which conflicts among older names, synonymies, and prevailing usage have required careful assessment under the Code.
The classification of all species within the genera and subgenera of the subfamily Polyommatinae, excluding the tribe Polyommatini, which will be treated in a future contribution, is critically reviewed and justified. Particular emphasis is placed on the generic classification of the tribe Scolitantidini, for which alternative taxonomic arrangements have been proposed and which remains a matter of taxonomic debate.

Key words
Taxonomy — Checklist — Papilionoidea — Lycaenidae — Polyommatinae — Polyommatini — Lampidini — Celastrinini — Scolitantidini — Everini — Zizeeriini — Azanini (tribe proposed) Danini — Azanina — Leptotes — Cyclyrius — Azanus — Lampides — Cacyreus — Celastrina — Tarucus — Phengaris — Maculinea — Pseudophengaris (genus proposed) —Turanana — Pseudophilotes — Palaeophilotes — Scolitantides — Iolana — Glaucopsyche — Zizeeria — Zizina — Cupido — Western Palearctic.

Addendum to section 1. Classification of tribes, subtribes, genera and subgenera within the subfamily Polyommatinae

Addendum to section 1.3. Classification of tribes, subtribes, genera and subgenera within the subfamily Polyommatinae – Conclusion

Two major publications dealing with the classification and nomenclature of the Polyommatinae were not discussed in Taymans & Cuvelier (2026, AWPL 2026(2): 27–35). These are Stradomsky (2016) and Zhang et al. (2023).
Stradomsky (2016) addressed the subject by considering morphological characters, most of which had already been studied by Eliot (1973), the variability of genital structures, and several genetic markers of the Polyommatinae. On this basis, the subfamily was divided into several tribes and subtribes, including several newly established taxa.
Based on a phylogenetic analysis, Zhang et al. (2023) proposed a comprehensive and detailed classification of the family-group taxa (subfamilies, tribes and subtribes) of the family Lycaenidae. The authors provided a summary table in which a substantial number of new taxa were introduced, particularly at the subtribal level. For the subfamily Polyommatinae, six tribes and 27 subtribes were recognized. In their classification, Polyommatini, the only tribe represented in the West Palaearctic region, comprises 23 subtribes. Apart from a few differences discussed below, this classification is broadly consistent with that of Stradomsky (2016).

Table 1. Comparison of the classifications of the subfamily Polyommatinae Swainson, 1827 proposed in the three aforementioned studies.

The differences between the two classifications are as follows:

• The tribes Oxylidini, Remelanini and Hypolycaenini were not studied by Stradomsky (2016).
• The arrangement of the subtribes within the tribe Polyommatini differs considerably between the two classifications (see table).
• The complex of subtribes Azanina/Unina/Danina/Ionolycina recognized by Zhang et al. (2023) comprises only two subtribes in Stradomsky (2016), namely Azanina and Danina.
• The genus Celastrina is included in the subtribe Lycaenopsina by Stradomsky (2016), whereas Zhang et al. (2023) place it in a separate subtribe, Celastrinina.
• The authorship of the family-group taxa is not given by Stradomsky (2016), whereas all taxa cited by Zhang et al. (2023) are followed by their authorship and date of publication. Approximately a dozen subtribal names are treated as new and are attributed to Grishin, one of the co-authors of the publication. Only Hemiolaina and Ionolycina had never previously been used and are therefore new taxa established by Grishin. Other subtribal taxa had already been named previously, but Zhang et al. (2023) considered them unavailable because their original authors had not explicitly indicated that the names were intentionally established as new. They were consequently (re)established by Grishin. These taxa are Cupidopsina, Azanina, Zizulina and Jamidina (previously mentioned by Koçak 1996), Unina, Fameganina and Pithecopina (previously mentioned by Koçak & Seven 1997), and Theclinesthina, Oboroniina and Uranothaumatina (previously mentioned by Stradomsky 2016).

The classification (Table 1) proposed in the present paper is based on the aforementioned publications. However, the tribe Polyommatini has been divided into several additional tribes in order to provide a more coherent structure and to better reflect the relationships among the different subtribes. This classification follows the monophyly of the various entities and is also consistent with the genital morphology of the genera included in the tribes recognized here. The table below provides a comparison of the three classifications discussed above.

Addendum to section 1.4. Proposed tribe Azanini

As the family-group taxon Azanina Grishin, 2023 was established after Danina Koçak & Seven, 1997, the latter name should be used for the tribe to which the genus Azanus belongs. The complete classification is therefore as follows:

Addendum to section 1.5. References

Koçak A. Ö. 1996. Güney batı asya Polyommatinae taksonlarının listesi ve bazı nomenklatür notları (Lepidoptera, Lycaenidae). — Centre for Entomological Studies Ankara Miscellaneous Papers 30/33(1–32). (url not available)

Koçak A. Ö. & Seven S. 1997. Negros Papilionoidea, Hesperioidea faunası ve taksonomisi üzerine araştırmalar (Lepidoptera) (Filipin entomolojik ekspedisyonu sonuçları-IV). — Priamus 9(2/3): 57-114. (url)

Stradomsky B.V. [Страдомский Б.В.]. 2016. A molecular phylogeny of the subfamily Polyommatinae (Lepidoptera: Lycaenidae) [Молекулярная филогения подсемейства Polyommatinae (Lepidoptera: Lycaenidae)]. — Кавказский энтомол. бюллетень [Caucasian Entomological Bulletin] 12(1): 145-156, fig. (url)

Taymans M. & Cuvelier S. 2026. Toward a revised checklist of the Western Palearctic butterflies, hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea) Part V: Rationale and framework for the Lycaenidae (part II) — AWPL 2026(2): 27–35. (url)

Zhang J., Cong Q., Shen J., Song L. & Grishin N.V. 2023. Butterfly classification and species discovery using genomics. — Taxonomic Report of the International Lepidoptera Survey 11(3): 1-94. (url)

Addendum to section 3. Pseudophilotes Beuret, 1958 and Palaeophilotes Forster, 1938 (Lycaenidae): nomenclatural and taxonomic considerations

3. Addendum 1. Recent developments

The recent publication by Lukhtanov et al. (2025), which had not been consulted prior to the publication of Taymans & Cuvelier (2026), substantially modifies the conclusions of the latter paper in two respects: the validity or otherwise of the taxon Palaeophilotes Forster, 1938, and the monophyly of the genera that were retained.

3. Addendum 2. Historical background and key facts

In 1938, Forster published the generic name Palaeophilotes as follows:
(original text in German):
Gattung: Palaeophilotes nov. gen.
triphysina Stgr.
Diese wenig bekannte zentralasiatische Art steht sowohl nach dem Habitus, wie auch nach dem Bau des ♂ Genitals völlig isoliert, sodaß ich mich gezwungen sehe, eine Gattung Palaeophilotes zu errichten. Palaeophilotes triphysina Stgr scheint ebenso wie Praephilotes anthracias Christ eine sehr primitive, altertümliche Art zu sein, was ja wohl schon daraus hervorgeht, daß bei keiner der beiden Arten blaue Färbung oder Androkonien ausgebildet sind.
(English translation):
Genus: Palaeophilotes nov. gen.
triphysina Stgr.
This little-known Central Asian species stands completely isolated both in its general appearance and in the structure of its male genitalia, so we feel compelled to establish a genus, Palaeophilotes. Palaeophilotes triphysina Stgr, like Praephilotes anthracias Christ, appears to be a very primitive and ancient species, as evidenced by the fact that neither species exhibits blue coloration or androconia.
Remark 1: Without referring to it in the preceding text, Forster published a plate (pl. 1, fig. 4) showing a photograph of the male genitalia of triphysina Stgr.
Remark 2: In the same paper, all other species traditionally associated with the genus Pseudophilotes were classified in the genus Philotes Scudder, 1876.

In 1958, Beuret published a thoroughly documented paper on the species of the tribe ‘Glaucopsychidi’ and described Pseudophilotes, gen. nov., to accommodate the following species: baton (type species), vicrama, abencerragus, bavius, and lanty, the latter species being currently classified in Glaucopsyche.

In 2013, Korb questioned the validity of the name Palaeophilotes, providing the following comment and establishing the replacement name Borisinia subgen. nov.:
(original text in Russian):
Данный диагноз не удовлетворяет требованиям Ст. 11 и 13.1 МКЗН [2004], так как не содержит собственно диагноза нового рода или его описания. В статье имеется изображение гениталий типового вида рода [Forster, 1938: Taf. I, Fig. 4], однако изображения не являются пригодными для установления названий после 1931 г. (Ст. 12, 13 МКЗН); кроме того, в
диагнозе рода нет указания на это изображение. Таким образом, название Palaeophilotes Forster, 1938 оказывается nomen nudum. Этот таксон следует переописать, что мы и делаем ниже.
(English translation):
This diagnosis does not satisfy the requirements of Articles 11 and 13.1 of the ICZN [2004], as it contains neither a diagnosis nor a description of the new genus. The paper includes an image of the genitalia of the type species of the genus [Forster, 1938: Taf. I, Fig. 4], but an illustration alone is not sufficient for the establishment of a name after 1931 (Articles 12 and 13 of the ICZN); moreover, the generic diagnosis does not refer to this image. Consequently, the name Palaeophilotes Forster, 1938 is a nomen nudum. This taxon should be redescribed, as is done below.

In 2022, Lukhtanov et al. published a paper on the molecular phylogeny of the subtribe Scolitantidina, confirming the validity of the genus Pseudophilotes and the division of its species into two subgenera, Pseudophilotes (for baton, vicrama, panoptes, sinaicus, jacuticus, barbagiae, abencerragus, panope) and Rubrapterus (for bavius, fatma). The species triphysina was not included in this study.

In 2023, Zhang et al. published a paper on the molecular phylogeny of a limited number of Lycaenidae species, but included triphysina. They concluded that this species is very closely related to three other species traditionally classified in Pseudophilotes and, consequently, that Palaeophilotes, being the older name, should also apply to the entire assemblage currently included in Pseudophilotes. Nevertheless, they retained the three subgenera Palaeophilotes, Pseudophilotes, and Rubrapterus, because these constituted monophyletic groups in their study.

In 2026, Taymans & Cuvelier, with the aim of promoting nomenclatural stability, raised each of the relevant taxa by one rank. Compared with Zhang et al. (2023), the genus Palaeophilotes was raised by one rank to become the subtribe Palaeophilotina, while the subgenera Palaeophilotes, Pseudophilotes, and Rubrapterus were likewise raised by one rank to become genera. This arrangement ensures that all species traditionally classified in Pseudophilotes remain within that lineage.

In 2025, Lukhtanov et al. published a paper on the molecular phylogeny of the genus Palaeophilotes and included all related taxa. They questioned Korb's (2013) interpretation and considered the taxon Palaeophilotes to be valid:
« As Article 13 of the Code states, names published after 1930 “must be accompanied by a description or definition that states in words characters that are purported to differentiate the taxon” (ICZN 1999). Such characters are indicated in the description. First, this is the structure of the genital apparatus of the males (“nach dem Bau des ♂ Genitals völlig isoliert”) (Forster 1938, page 105). Additionally, Forster states in words that the only species of the genus Palaeophilotes (Palaeophilotes triphysina) and the only species of the genus Praephilotes (Praephilotes anthracias) share such unique characters as the absence of blue color and androconia on the wings. Then, on the page 106 of the same article, Forster notes the unique feature of Praephilotes anthracias (“Der viel gestrecktere Flügelschnitt,” thus completing the diagnosis of and the formal description to satisfy Art. 13.
Thus, considering the availability of the name Palaeophilotes, the name Borisinia is a junior objective synonym of Palaeophilotes, since it is based on the same type species. »

Furthermore, they demonstrated that triphysina (the type species of the genus Palaeophilotes) is nested within the species traditionally classified in Pseudophilotes. The option of retaining several subgenera, as proposed by Zhang et al. (2023), is therefore no longer tenable because these subgenera are no longer monophyletic (with the exception of the bavius/fatma lineage, which remains monophyletic). They consequently placed all species in Palaeophilotes, without recognising any subgenera.

3. Addendum 3. Validity of the taxon Palaeophilotes Forster, 1938

Article 13.1 of the ICZN Code stipulates that, for a taxon of generic rank published after 1930 to be valid, the author must provide ‘a description or definition that states in words characters that are purported to differentiate the taxon’.
Is there a description that allows the taxon to be differentiated? No. Is there a definition that allows the taxon to be differentiated? There is a definition, but does it allow the taxon to be differentiated?
Forster's 1938 publication contains two definitions. The first, ‘completely isolated both in its general appearance and in the structure of its male genitalia’, could apply to a considerable number species of butterflies. The second, ‘Palaeophilotes triphysina Stgr, like Praephilotes anthracias Christ, appears to be a very primitive, ancient species, as evidenced by the fact that neither species exhibits blue coloration or androconia’, does not provide a diagnostic character because the distinction invoked applies to both taxa mentioned and therefore does not allow them to be differentiated.
From a purely textual perspective, the fact that a definition is provided could be interpreted in several ways, and the arguments put forward by Korb (2013) or Lukhtanov et al. (2025) could therefore be considered valid. However, the fact remains that the publication contains no element that provides a genuine diagnosis. We therefore retain the interpretation of Korb (2013) in our classification.

3. Addendum 4. Generic structure to be adopted in the classification

The study by Lukhtanov et al. (2025) demonstrates that the relationships among the different species follow the pattern below (available genera for the species concerned are given in parentheses):

To satisfy the requirement that generic entities be monophyletic, the following classification is proposed:
Genus Pseudophilotes Beuret, 1958
(TS: Papilio baton Bergsträsser, [1779])
Subgenus Pseudophilotes (Pseudophilotes) Beuret, 1958
(= Palaeophilotes Forster, 1938; = Inderskia Korshunov, 2000; = Borisinia Korb, 2013)
P. (P.) abencerragus (Pierret, 1837)
P. (P.) barbagiae (De Prins & Poorten, 1982)
P. (P.) panoptes (Hübner, [1813])
P. (P.) baton (Bergsträsser, [1779])
P. (P.) sinaicus (Nakamura, 1975)
P. (P.) vicrama (Moore, 1865)
P. (P.) jacuticus (Korshunov & Viidalepp, 1980)
P. (P.) panope (Eversmann, 1851)
P. (P.) triphysina (Staudinger, [1892])
Subgenus Pseudophilotes (Rubrapterus) Korshunov, 1987
P. (R.) bavius (Eversmann, 1832)
P. (R.) fatma (Oberthür, 1890)

3. Addendum 5. References

Beuret H. 1958. Zur systematischen Stellung einiger wenig bekannter Glaucopsychidi (Lep., Lycaenidae). — Mitteilungen der entomologischen Gesellschaft Basel NF 8(6): 61-79 (url), 81-100 (url).

Forster W. 1938. Das System der paläarktischen Polyommatini. (Lep. Lycaen.). — Mitteilungen der Münchner Entomologischen Gesellschaft (e.V.) 28(2): 97-118, pl. 1-3. (url)

Корб С.К. [Korb S.K.]. 2013. К систематике, биологии и диагностике малоизвестного вида, описанного как Lycaena triphysina Staudinger, 1891 (Lepidoptera: Lycaenidae) [To the systematics, biology and diagnostics of a little known species Lycaena triphysina Staudinger, 1891 (Lepidoptera: Lycaenidae)]. — Эверсманния. ЭнтомологическиеисследованиявРоссииисоседнихрегионах [Eversmannia] 34: 32-37. (url)

Lukhtanov V. A., Makhov I. A., Gagarina A. V. & Romanovich A. 2025, Taxonomy of the West Palaearctic butterfly genus Palaeophilotes Forster, 1938 (Lepidoptera: Lycaenidae) based on combined analysis of COI barcodes and multilocus nuclear markers. — Zootaxa 5584(4): 570-580. https://doi.org/10.11646/zootaxa.5584.4.8

Taymans M. & Cuvelier S. 2026. Toward a revised checklist of the Western Palearctic butterflies, hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea) Part V: Rationale and framework for the Lycaenidae (part II) — AWPL 2026(2): 27–35. (url)

Author contribution
Michel Taymans: conceptualisation, analysis, visualisation, writing - original draft, writing – review and editing.
Sylvain Cuvelier: analysis, validation, visualisation, writing – review and editing.

Acknowledgements
We are sincerely grateful to Theo Garrevoet for his careful and thorough review of the final draft, and we wish to acknowledge Bénédicte Jonckers for her invaluable administrative support as production editor.