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

. This paper discusses three problems concerning the Woodland Grayling, Hipparchia fagi Scopoli, 1763, with respect to the identity and application of the junior name Papilio hermione Linnaeus, 1764. In 1977, the late Otakar Kudrna designated a specimen of the Rock Grayling, Hipparchia alcyone [Denis & Schiffermüller], 1775, to become the lectotype of Papilio hermione – as a result of which hermione supplant-ed alcyone as the senior epithet for this species. Because P. hermione is the nominal type species of Hipparchia Fabricius, 1807, Kudrna’s action rendered this a genus based on a misidentified species. Third, while a majority of lepidopterists have ignored Kudrna’s action and continue to apply the name H. alcyone to the Rock Grayling, and still regard P. hermione as a junior subjective synonym of H. fagi , the formal nomenclature for the Rock Grayling has become unstable because a large minority have nonetheless accepted Kudrna’s lectotype designation and all that follows from it. It is demonstrated here that no syntypes of Papilio hermione (or Papilio fagi ) have survived; consequently, Kudrna’s lectotype designation for P. hermione is invalid. By designation of a single specimen of the Woodland Grayling as neotype for both P. fagi and P. hermione , the two names are rendered objectively synonymous, thereby restoring stability to the species name for the Rock Grayling (as Hipparchia alcyone ), and to the application of Papilio hermione (= Hipparchia fagi ) as nominal type species of the generic name Hipparchia .


Introduction
The history of Papilio hermione Linnaeus, 1764 The name Papilio hermione was based on a specimen or specimens in the collection of Queen Louisa Ulrika (1720-1782) of Sweden, who had asked Linnaeus to catalogue her collections housed at Drottningholm Palace; the name was published by Linnaeus in 1764 (Fig. 1). Three years later, Linnaeus (1767: 773) listed Papilio fagi Scopoli, 1763 (Fig. 2) as a synonym of P. hermione. Over the following two centuries, the synonymy of these two names for the Woodland Grayling was accepted by most workers. However, some (e.g. Fabricius 1793: 232/3, and Turton 1806: 32) continued to use Linnaeus's name without any reference to fagi [Note however that Aurivillius (1882: 81/82) suggested that Turton's usage represents a misidentification of Chazara briseis (Linnaeus, 1764)]. Others (e.g. Doubleday 1845: 130;Kirby 1871: 86, 705), while explicitly accepting the synonymy, were slow to adopt Scopoli's fagi as the senior name. At that time, priority was less rigidly applied than today. Recognising that there were two rather similar grayling species, the Woodland Grayling and the usually smaller Rock Grayling, [Denis and Schiffermüller] (1775), acting as first revisers (ICZN 1999: Article 24.2), accepted that P. fagi was applicable to the larger species and proposed the name Papilio alcyone for the smaller Rock Grayling. Although the description of P. fagi by Scopoli is not definitive (Verity 1913: 158), his associated figure (Fig. 3) is clearly discernible as the larger Woodland Grayling. This nomenclatural synonymy remained stable until Hoffmansegg (1804) realised that Cramer (1775) had used the specific epithet Papilio alcyone for a completely different butterfly, the pierid currently known as Catopsilia pyranthe (Linnaeus, 1758).
This discovery was largely ignored at first, the nomenclatural synonymy remaining stable, until Hemming (1943) highlighted this problem and suggested that Papilio aelia Hoffmansegg, 1804, should replace alcyone [Denis & Schiffermüller] as the name for the Rock Grayling. This was accepted by Verity (1953) and Forster and Wohlfahrt (1955). However, for the sake of nomenclatural stability, the International Commission for Zoological Nomenclature (ICZN) subsequently decided that the publication of [Denis and Schiffermüller] (1775) should be considered to predate that of Cramer (1775) (Opinion 516, 1958); thus, stability was restored and Hipparchia alcyone remained the accepted name for the Rock Grayling.
Stability was, however, once again upset by the designation of a lectotype for the name Papilio hermione by Kudrna (1977: 27) in his revision of the genus Hipparchia. Unfortunately, Kudrna decided to go against the then prevailing usage and designated a specimen of H. alcyone with a label 'hermione' on the pin in Linnaeus's handwriting. This designation was both unnecessary and invalid, as hermione was not then in use, and the specimen designated was not a syntype -as it has to be under the 'code' (ICZN 1999: Article 74.2). When this designation was challenged by Higgins and Riley (1978), it was suggested by Kudrna (1984) that the specimen could have been removed by Linnaeus from Queen Ulrika's collection (which would place it as a syntype) and added it to his own. There is no proof whatsoever for this contentious statement. A label on a pin has no nomenclatural status, as it is well-known and appreciated that labels can be moved, removed, acquired and placed on the pins of other specimens (Brown 1968: 85;Honey and Scoble 2001: 331). The suggestion by the late Ahmet Ömer Koçak (1983: 168) that the name hermione Linnaeus, 1764, should be treated as a nomen dubium, because of the apparent lack of any syntypes, which in some ways would seem very reasonable, was not accepted by Kudrna (1984: 234).
What species could Queen Ulrika's P. hermione specimens have represented? Brown (1968: 77) stated that Carl Clerck (1759 & 1764) was commissioned by Linnaeus to paint the butterflies in Queen Ulrika's collections for the sake of posterity, knowing the fragility of butterflies. The two volumes of published paintings were examined but no painting of Papilio hermione was found. Brown also stated that there were some unpublished and unbound paintings held by the Library of the Swedish Academy of Sciences in Stockholm; this was confirmed by Maria Dahlbäck (see acknowledgements). The first author visited the Library and examined these paintings; however, unfortunately, they did not yield a painting of the butterfly named hermione either.
Whilst in Sweden, the remnants of the Queen's collection in Uppsala were examined; there was definitely no specimen of hermione extant, consistent with the observations of Carl Peter Thunberg (1743-1828), who listed the specimens in the Queen's collection when it arrived at Uppsala in 1803 ( Fig. 4; see also Aurivillius 1882). The entomological collections of Uppsala University also include Thunberg's own collection, in which he used the names of his mentor (Linnaeus), rather than possibly more appropriate (i.e. earlier) names. Four butterflies included under 'hermione' (Fig. 5;3♂♂,1♀) are all considered to be Hipparchia fagi originating from old German populations (D. Jutzeler, pers. comm. to first author); Linnaeus (1764: 281) gave 'Germania' as the 'habitat' [type locality] for Papilio hermione.
On this slender basis it would seem more likely that Queen Ulrika's specimen or specimens of P. hermione represented the species known today as Papilio fagi Scopoli, rather than any other grayling butterfly.

Why Kudrna's lectotype designation is invalid
The following points relate to the lectotype designation of Kudrna (1977: 27): • Linnaeus (1764) based the name Papilio hermione on a butterfly specimen or specimens present in the collection of Queen Ulrika of Sweden (see above). • Linnaeus also cited illustrations by James Petiver (1702, p. 12, pl. 7, fig. 5) and August Johann Rösel von Rosenhof (1761, pl. 27, figs 3, 4; Fig. 1). According to Honey and Scoble (2001: 331), these represent Hipparchia fidia (Linnaeus, 1767) (currently known as Pseudotergumia fidia) and Brintesia circe (Fabricius, 1775), respectively. According to John Chainey (MS), the Petiver material no longer exists, while, according to Horn and Kahle (1936, p. 228), the insect collection of Rösel von Rosenhof is missing ("verschollen"). • Queen Ulrika's collection fell into neglect following her death in 1782 (Honey and Scoble 2001: 285). • The Queen's collection was bequeathed by King Gustav to Uppsala University in 1803 and was catalogued by Carl Peter Thunberg (one of Linnaeus's 'Apostles' -well-thought of students) on its arrival in Uppsala in 1804, by which time Linnaeus was dead (deceased 1778). • There was no specimen of P. hermione present in the Queen's collection in 1804 (see Fig. 4) and any original specimens must have been a casualty of the aforementioned neglect (see also above). • There is no credible evidence to suggest that any of the Hipparchia specimens in the Linnaean Society of London collection could have come from Queen Ulrika's collection (see above). • In conclusion, there are no syntypes available for selection of a lectotype for the name Papilio hermione Linnaeus; consequently, Kudrna's (1977) designation is invalid. Fabricius (1807, pp. 281/2) evidently included 119 species in his new genus Hipparchia, explicitly listing nine of these but without indicating a type: Papilio hermione, P. fauna, P. maera, P. ligea, P. epiphron, P. galathea, P. pilosellae, P. hyperanthus, and P. rumina. With the exception of the last (a papilionid), all represent members of the Nymphalidae: Satyrinae, although only two are now included in the genus Hipparchia as currently circumscribed: P. hermione and P. fauna. The first person to make a valid designation of one of these names to be the type species of Hipparchia was Butler (1868b, p. 50), who stated "Typical Species Hipparchia Fagi (Hermione)." As Papilio fagi was not included in Fabricius's list, the only possible interpretation of Butler's cryptic statement is that he selected Papilio hermione -but in the firm belief that it represented the same taxon as the senior name Papilio fagi Scopoli, based no doubt and, as presented above, on Linnaeus's (1767) own synonymy.

A second problem created by Kudrna's lectotype designation
An issue affecting this type species designation was clarified by Cowan (1985), who realised that Butler published two papers in the same year that dealt with Hipparchia: one in February (Butler 1868a) and the second in July (Butler 1868b). Hemming (1967: 220), while correctly indicating that Butler had selected Papilio hermione as type species, incorrectly cited the February paper, not the later work. Koçak (1983: 167) and Kudrna (1984: 229) were thus misled or failed to appreciate that Butler's later publication was the relevant source. As a result, because the first of these papers (Butler 1868a: 194) did not mention or associate P. hermione with Hipparchia fagi, they rejected Butler's designation, on the grounds that Papilio fagi Scopoli, 1763, was not one of the species listed by Fabricius (1807). However, as Cowan made clear, and as indicated above, Butler's (1868b) later selection of 'Hipparchia Fagi (Hermione)' is valid. Kudrna's (1977) action, because it broke the synonymy of P. fagi and P. hermione accepted by Linnaeus (1767) and, critically, Butler (1868b), rendered Hipparchia a genus based on a misidentified type species (ICZN 1999, Article 70.3). Rejecting Kudrna's action, together with selection of a single neotype to make P. hermione and P. fagi objective synonyms (see below), preserves application of the genus Hipparchia -as the nominal type species, Papilio hermione Linnaeus, is thus now objectively the same taxon as represented by the earlier-established nominal species, Papilio fagi Scopoli.

Scopoli's butterflies
None of Scopoli's collection of butterflies is known to have survived; they were almost certainly all lost when his house in Idrija caught fire. A statement by Soban (2005) on the life of Scopoli reports as follows: "Similar misfortunes accompanied him in Idrija where he was to remain for 15 years. He again lost everything in a fire which destroyed his home and all of his family." Thus there are no syntypes of Papilio fagi either.

Neotype designation
As there is no surviving type material for the nominal taxa Papilio hermione and P. fagi, we hereby designate a single specimen of Hipparchia fagi (Scopoli, 1763) as neotype for both. The specimen originates from the Austrian state of Styria, a general region where the original material of P. hermione ('Germania') and P. fagi (Carniola = Krain) could well have come from. In 18 th century Europe, Germany consisted of multiple 'Electorates' and 'Kingdoms', including the Archduchy of Austria. Carniola, itself formerly part of the Archduchy, comprised most of present day Slovenia and small areas of Croatia and Italy, and extended north-east as far as Styria, in the southern border region of modern Austria.
The neotype is a fresh male from Kreuzberg (Styria = Steiermark), showing the wing characters typical of the species almost universally known by the name Hipparchia fagi. Deposited in the Tiroler Landesmuseen, Innsbruck, Austria (TLMF), the labels on the pin are as follows: on off-white paper Kreuzberg is located about 6 km north of the Slovenian border, and 3 km north of Leutschach in southern Austria, where it rises to 633 m. Kleingraben refers to a valley or small river. The specimen is figured upperside and underside, together with the labels on the pin (Fig. 6A-C, respectively). A specimen of this species was chosen for two reasons: firstly Linnaeus himself made the synonymy

C.
in 1767 and, secondly, because this action will reinstate nomenclatural stability by making P. hermione an objective junior synonym of H. fagi, a synonymy that was almost consistently accepted for over 200 years until destabilised by Kudrna (1977).

A third need for re-establishing the synonymy
Further evidence for the necessity to re-establish this synonymy and thus stability can be seen from the appendices, which include: first, a list of 62 works (including a website) by authors who ignored Kudrna's (invalid) lectotype designation and placed H. hermione as a synonym of H. fagi (Appendix 1) and, secondly, 26 publications and websites that accepted the lectotype designation by Kudrna (1977) and placed H. alcyone as a synonym of H. hermione (Appendix 2).
A further consequence of this neotype designation is that the senior name for the Rock Grayling will be re-established as Hipparchia alcyone [Denis & Schiffermüller, 1775], a name which a majority of authors still adopt today in preference to 'hermione' sensu Kudrna (see Appendix 1). Note also, however, the issue of separation of Hipparchia genava (Fruhstorfer, 1908) from H. alcyone at species level