Butterfly Name Restored After 200-Year Taxonomic Dispute

For more than two centuries, the scientific community has relied on a stable nomenclature for the Woodland Grayling butterfly, known formally as Hipparchia fagi. This stability was rooted in the work of Carl Linnaeus, who in 1767 explicitly listed Papilio fagi as a synonym of Papilio hermione, a name he had published the previous year. For two hundred years, lepidopterists accepted this synonymy, treating the two names as interchangeable references to the same species. However, a single taxonomic action taken in 1977 by the late Otakar Kudrna disrupted this long-standing consensus, creating a complex web of nomenclatural instability that has persisted for nearly five decades. The recent publication in Nota Lepidopterologica addresses this issue directly, demonstrating that Kudrna’s designation was invalid and proposing a solution that restores the original stability of these names.

The controversy centers on the identity of the type species for the genus Hipparchia, which was established by Fabricius in 1807. In 1868, the entomologist Walter Rothschild’s contemporary, A. E. Butler, designated Papilio hermione as the nominal type species for this genus. Butler made this choice under the firm belief that hermione was a subjective synonym of the senior name, Papilio fagi. This designation was accepted by the scientific community because it aligned with Linnaeus’s own earlier synonymy. The situation remained stable until Kudrna, in his 1977 revision of the genus, designated a specific specimen of the Rock Grayling, Hipparchia alcyone, as the lectotype for Papilio hermione. This action effectively broke the synonymy between hermione and fagi, rendering the genus Hipparchia based on a misidentified species and causing significant confusion in the application of these names.

The validity of Kudrna’s lectotype designation has been challenged on several grounds, primarily concerning the survival of the original type material. Linnaeus based the name Papilio hermione on specimens from the collection of Queen Louisa Ulrika of Sweden. Historical records indicate that this collection fell into neglect after the Queen’s death in 1782. When the collection was eventually bequeathed to Uppsala University in 1803 and catalogued by Carl Peter Thunberg, no specimen of Papilio hermione was present. This absence suggests that any original syntypes had been lost to neglect or decay before the collection reached the university. The authors of the recent paper examined the remnants of the Queen’s collection in Uppsala and confirmed that no specimen of hermione extant, consistent with Thunberg’s earlier observations.

Further investigation into the historical context of Linnaeus’s naming reveals that he likely muddled together multiple species of graylings. Linnaeus cited illustrations by James Petiver and August Johann Rösel von Rosenhof in his 1764 publication. Modern analysis suggests that these illustrations represent different species, specifically Hipparchia fidia and Brintesia circe. This indicates that Linnaeus’s original concept of Papilio hermione was not based on a single, clearly defined taxon. The lack of surviving syntypes from the Queen’s collection, combined with the ambiguity of the original illustrations, means that there is no valid material from which a lectotype could be selected. Therefore, Kudrna’s 1977 designation, which relied on a specimen that was not a syntype, is technically invalid under the rules of the International Commission on Zoological Nomenclature.

The consequences of Kudrna’s invalid designation have been far-reaching, leading to a split in the scientific community regarding the correct name for the Rock Grayling. A majority of lepidopterists have ignored Kudrna’s action and continue to apply the name Hipparchia alcyone to the Rock Grayling, maintaining the traditional synonymy of hermione with fagi. However, a significant minority have accepted Kudrna’s lectotype designation, leading to the application of the name hermione to the Rock Grayling and the designation of alcyone as a junior synonym. This divergence has created nomenclatural instability, with different publications and databases using different names for the same species, complicating research and conservation efforts.

To resolve this issue, the authors propose the designation of a neotype for both Papilio hermione and Papilio fagi. A neotype is a specimen selected to serve as the single physical embodiment of the type status of a nominal taxon when the original type material is lost or destroyed. By designating a single specimen of the Woodland Grayling as the neotype for both names, the authors render the two names objectively synonymous. This action restores the synonymy that was accepted by Linnaeus and Butler, thereby stabilizing the application of the name Papilio hermione as the nominal type species of the genus Hipparchia. The neotype specimen is a fresh male from Kreuzberg in Styria, Austria, collected in 1979 and deposited in the Tiroler Landesmuseen in Innsbruck.

The choice of the neotype specimen was guided by historical and geographical considerations. Kreuzberg is located in a region that corresponds to the historical areas of Germania and Carniola, where the original material for Papilio hermione and Papilio fagi likely originated. In the 18th century, these regions were part of the broader German-speaking lands, including the Archduchy of Austria. The specimen exhibits the wing characters typical of the species universally known as Hipparchia fagi, ensuring that the neotype accurately represents the taxon intended by Linnaeus and Scopoli. This designation provides a clear and objective reference point for future taxonomic work, eliminating the ambiguity that has plagued the nomenclature of these butterflies for decades.

The restoration of the synonymy between Papilio hermione and Papilio fagi has significant implications for the stability of the genus Hipparchia. By ensuring that the nominal type species is correctly identified as the Woodland Grayling, the authors preserve the application of the genus name as originally intended by Fabricius and Butler. This action also re-establishes the senior name for the Rock Grayling as Hipparchia alcyone, a name that has been widely used by the majority of authors. The resolution of this taxonomic dispute not only clarifies the nomenclature of these specific species but also serves as a case study in the importance of adhering to the rules of zoological nomenclature and the need for careful examination of historical type material.

The recent publication in Nota Lepidopterologica represents a significant contribution to the field of lepidopterology, providing a clear and well-supported solution to a long-standing nomenclatural problem. By demonstrating the invalidity of Kudrna’s lectotype designation and proposing a valid neotype, the authors restore stability to the application of the names Papilio hermione and Papilio fagi. This work underscores the importance of historical context and rigorous taxonomic analysis in resolving disputes over scientific names. As the scientific community continues to refine its understanding of biodiversity, such efforts are essential for maintaining the accuracy and consistency of biological nomenclature.

Subject of Research: Taxonomic nomenclature and type species designation in Satyrinae butterflies

Article Title:  Papilio hermione Linnaeus, type species of Hipparchia Fabricius ( Lepidoptera , Satyrinae ): restoring stability to the application of these names

Article References: Russell, P. J. C., & Vane-Wright, R. I. (2022). Papilio hermione Linnaeus, type species of Hipparchia Fabricius (Lepidoptera, Satyrinae): restoring stability to the application of these names. Nota Lepidopterologica, 45, 279-294. https://doi.org/10.3897/nl.45.85341

Image Credits: AI Generated

DOI: 10.3897/nl.45.85341

Keywords: Lepidoptera, Taxonomy, Nomenclature, Hipparchia, Neotype, Lectotype, Butterflies, Zoological Nomenclature, Systematics, Entomology, Papilio, hermione