Dynamic Checklist of the Western Palearctic butterflies
(Lepidoptera, Papilionoidea) Hyperlinked to Systematics, Genetics and Distribution Papilionoidea Rationale
The group-family level classification adopted in the checklist (Taymans & Cuvelier 2025), encompassing superfamilies, families, subfamilies, tribes and subtribes, largely follows the most recent phylogenetic studies on Pieridae (Braby et al. 2006; Wahlberg et al. 2014; Ding & Zhang 2016; Wei et al., 2022; Kawahara et al., 2023; Carvalho et al. 2024), with the notable exception of the Aporiini. The rationale for this exception and the approach adopted are discussed below.
Context
Unfortunately, the International Code of Zoological Nomenclature (ICZN) provides no biological, morphological, phylogenetic or genetic criteria for the recognition of taxa at the group-family level. The Code regulates nomenclature rather than taxa themselves and therefore does not define what constitutes a family or how it should be delineated.
Prior to phylogenetic analyses, group-family ranks were assigned at the discretion of individual systematists, based on a variety of criteria. These could include the structure of the exoskeleton, wing venation, legs, palpi, genitalia or life-history traits such as eggs, larvae and pupae. By considering these characteristics, systematists aimed to provide a conceptual framework for grouping species progressively into increasingly smaller sets, ultimately resulting in genus-level classifications that comprised only a few species.
In contrast, contemporary phylogenetic studies adopt a more objective approach by linking taxonomic ranks to the historical timing of lineage divergences. For instance, in Kawahara et al. (2023), the temporal intervals used to define classification levels are: families between 100 and 80 Ma, subfamilies between 80 and 60 Ma, tribes between 60 and 40 Ma, and subtribes between 40 and 20 Ma. This method is therefore anchored in estimated divergence times derived from mathematical models and can be considered less subjectively influenced. These intervals do not necessarily correspond to geological epochs (e.g., the 60 to 40 Ma interval used to define tribes spans the mid-Palaeocene to mid-Eocene). Nevertheless, the advantage of this approach is that it bases classification on a consistent, objective criterion, whereas its drawback is the exclusion of other potentially informative characteristics.
Conclusion
Regarding the Aporiini/Aporiina, Kawahara et al. (2023) recognise the group’s monophyly but include it within the Pierini. Their supplementary phylogenetic tree indicates that the Aporiini/Aporiina branch diverged from the Pierini at an estimated 36 Ma. Thus, based on an estimated 4 Ma divergence interval (subjectively calculated), the Aporiini/Aporiina are treated as a subtribe.
The last author to consider Aporiini/Aporiina at the tribe level was Braby (2005) in his Provisional Checklist of Genera of the Pieridae. What is the rationale for following this relatively older study rather than more recent ones? First, from a technical perspective, available evidence strongly supports the monophyly of the group. Second, although it is represented by a single species in the European fauna, the group is widely distributed globally, encompassing over 400 species. Recognizing it as a tribe allows for further subdivision into subtribes; for example, Belenois (and closely related genera) could constitute a distinct subtribe, separate from Delias (and related genera) or other genus-level clusters.
Braby M., Vila R. & Pierce N. 2006. Molecular phylogeny and systematics of the Pieridae (Lepidoptera: Papilionoidea): higher classification and biogeography. — Zoological Journal of the Linnean Society 147: 239–275. https://doi.org/10.1111/J.1096-3642.2006.00218.X
Carvalho A., Owens H, St Laurent R., Earl C., Dexter K., Messcher R., Willmott K., Aduse-Poku K., Collins C., Homziak N., Hoshizaki S., Hsu Y., Kizhakke A., Kunte K., Martins D., Mega N. Morinaka S., Peggie D., Romanowski H., Sáfian S., Vila R., Wang H., Braby M., Espeland M., Breinholt J., Pierce N., Kawahara A., Lohman D. 2024. Comprehensive phylogeny of Pieridae butterflies reveals strong correlation between diversification and temperature. — iScience 27(4): (109336): 1-13. https://doi.org/10.1016/j.isci.2024.109336
Ding C. & Zhang Y. 2016. Phylogenetic relationships of Pieridae (Lepidoptera: Papilionoidea) in China based on seven gene fragments. — Entomological Science 20(1): 15-23. https://doi.org/10.1111/ens.12214
Kawahara A., Storer C., Carvalho A., Plotkin D., Condamine F., Braga M., Ellis E., St Laurent R., Li X., Barve V., Cai L., Earl C., Frandsen B., Owens H., Valencia-Montoya W., Aduse-Poku K., Toussaint E., Dexter K., Doleck T., Markee A., Messcher R., Nguyen Y., Badon J., BenĂtez H., Braby M., Buenavente P., Chan W., Collins S., Rabideau Childers R., Dankowicz E., Eastwood R., Fric Z., Gott R., Hall J., Hallwachs W., Hardy N., Hawkins Sipe R., Heath A., Hinolan J., Homziak N., Hsu Y., Inayoshi Y, Itliong M., Janzen D., Kitching I., Kunte K., Lamas G., Landis M., Larsen E., Larsen T., Leong J., Lukhtanov V., Maier C., Martinez J., Martins D., Maruyama K., Maunsell S., Mega N., Monastyrskii A., Morais A., MĂĽller C., Naive M., Nielsen G., PadrĂłn P., Peggie D., Romanowski H., Sáfián S., Saito M., Schröder S., Shirey V., Soltis D., Soltis P., Sourakov A., Talavera G., Vila R., Vlasanek P., Wang H., Warren A., Willmott K., Yago M., Jetz W., Jarzyna M., Breinholt J., Espeland M., Ries L., Guralnick R., Pierce N. & Lohman D. 2023. A global phylogeny of butterflies reveals their evolutionary history, ancestral hosts and biogeographic origins. — Nature Ecology & Evolution 7: 903-913. Article: https://doi.org/10.1038/s41559-023-02041-9 . Supplementary Materials: url.
Taymans M. & Cuvelier S. A dynamic checklist of the Western Palearctic butterflies hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea). — Archives of Western Palearctic Lepidoptera 2025(1): 1-70. https://doi.org/10.5281/zenodo.14733224
Wahlberg N., Rota J., Braby M., Pierce N. & Wheat C. 2014. Revised systematics and higher classification of pierid butterflies (Lepidoptera: Pieridae) based on molecular data. — Zoologica Scripta, 43, 641–650. https://doi.org/10.1111/zsc.12075.
Wei F., Huang W., Fang L., He B., Zhao Y., Zhang Y., Shu Z., Su C. & Hao J. 2023. Spatio-Temporal Evolutionary Patterns of the Pieridae Butterflies (Lepidoptera: Papilionoidea) Inferred from Mitogenomic Data. — Genes 14(1):72. Article: https://doi.org/10.3390/genes14010072. Supplementary Materials: url.
Archives of Western Palearctic Lepidoptera Editors-in-Chief: Michel Taymans & Sylvain Cuvelier
