Satyrus cantheus Godart, [1824]

Cong, Qian, Barbosa, Eduardo P., Marín, Mario A., Freitas, André V. L., Lamas, Gerardo & Grishin, Nick V., 2021, Two new species of Hermeuptychia from North America and three neotype designations (Nymphalidae: Satyrinae), The Taxonomic Report of the International Lepidoptera Survey 9 (7), pp. 1-21 : 13-16

Satyrus cantheus Godart, [1824] is a junior subjective synonym of Hermeuptychia sosybius ( Fabricius, 1793)

As detailed by Cardé et al. (1970) and further elaborated by Viloria (2021), Godart ([1824]) divided the Fabrician concept of Papilio canthus Linnaeus, 1767 into three species. Two of them are known today as Lethe eurydice (Linnaeus, 1763) (type locality USA: Pennsylvania) and Emeryus argulus (Godart, [1824]) (type locality Brazil: Pará, Santa Bárbara do Pará). The latter name was proposed in the same work that also coined the name for the third species: Satyrus cantheus Godart, [1824] , which is a nomen dubium ( Cardé et al. 1970). Viloria (2021) suggested that one of the two specimens in the Hunterian Collection in Glasgow pinned below the header label " Pap. Canthus / Fabr. pag 64 N o 288" may have been used as a basis for the description of S. cantheus . Viloria discussed only the second specimen (GLAHM:127581, which is a female, not a male), without mentioning the first one (GLAHM: 127580, a male). These two specimens are of different species that are presently placed in different subtribes. The phenotype of this second specimen discussed by Viloria (GLAHM:127581) does not agree with the original description of S. cantheus . First, each of its forewings has five obvious eyespots instead of "three small eyespots not very pronounced" per the description. It is apparent, that Godart paid much attention to the number of eyespots and their structure. E. g., he separated argulus from canthus with cantheus (all of which Fabricius combined under P. canthus Linnaeus, 1767 ) by the number of ventral hindwing eyespots: 5 in the former versus 6 in the two latter species; and their structure: with two (not one) pupils in the former species. Second, the 5 th eyespot (next to the 6 th, which is stated to be at the anal angle [=tornus]) is not the largest: it is 1.2–1.3 times smaller than the 2 nd eyespot, measured in longitudinal dimension. Third, the 6 th eyespot (the one stated to be by the anal angle) is not the smallest: it is about 1.3 times larger than the 4 th one. Fourth, the specimen is about 1.5 times smaller than a typical L. eurydice , but Godart's description states that S. cantheus [Godart species number 56] "is about the same in size as the previous one", which is the number 55 P. canthus Linnaeus, 1767 (a junior objective synonym of L. eurydice ). For all these reasons, this specimen is not a syntype. Furthermore, this specimen would not define the name S. cantheus as intended in the original description to qualify as a neotype.

Fabricius did not propose a new name canthus homonymous with Papilio canthus Linnaeus, 1767 as suggested by Viloria (2021). This is because Fabricius gave the name canthus in the third line of his description and attributed it to the Linnaeus citation, in a manner he did for many other species when giving brief modified descriptions for previously proposed names ( Fabricius 1775). Moreover, he repeated this treatment and attribution in subsequent publications, only adding an additional name and citation to it ( Papilio argante Cramer, 1779 ) that he considered to refer to the same species ( Fabricius 1781 -[1782]; Fabricius 1793). No new name currently attributed to Fabricius that we know of referenced an identically spelled name citing a previous author and publication. Therefore, Fabricius either misidentified the Linnaean taxon completely, or expanded it to cover additional phenotypes, and Godart's work points to the latter ( Godart [1824]). Moreover, all new names proposed by Fabricius have been thoroughly studied and catalogued in detail by a number of authors, and none mentions canthus as a new name. For instance, there is no mention of canthus with Fabricius as the author in the comprehensive catalogue by Zimsen (1964). There is no name canthus assigned to Fabricius in The Global Lepidoptera Names Index ( Beccaloni et al. 2003). Furthermore, Godart ([1824]: 493) did not credit canthus to Fabricius, only to Linnaeus, mentioning that "Fabricius combined this butterfly [i.e. canthus ] with the next [called cantheus by Godart]; but it appears to us to be a different species, at least judging from the description that follows" (interpretively translated and comments in brackets are added by us). Because Fabricius did not propose a new name canthus , no name-bearing (i.e., type) specimens of the Fabrician concept of canthus are possible by definition: there was no new name to bear.

We agree with Cardé et al. (1970) and Viloria (2021) that Godart's S. cantheus is not a misspelling of P. canthus , but is an available name. Our reading of Godart ([1824]) excludes the possibility of a misspelling, because Godart uses " S. canthus " for his species no. 55 and " S. cantheus " for his species no. 56: the two consecutively numbered species that are expected to have different names, as they do (not one being a misspelling of the other). Miller and Brown (1981) made a mistake in stating that Godart attributed the name cantheus to Linnaeus: only the name canthus [no. 55] was attributed to Linnaeus, not the name number 56, which cited only Fabricius. Therefore, the conclusion of Miller and Brown about the misspelling is erroneous. Furthermore, Godart used two sets of names throughout his work: French names that were not italicized, employed French words and had accents over letters when appropriate; and Latin names that were italicized, employed Latin or Latinized words and lacked accented letters. For instance, he used "Satyre" as a French name, which would be "Satyr" in English, not the Latin Satyrus , and therefore not a genus epithet. With this distinction, only Godart's Latin names would enter zoological nomenclature. For instance, for the species known today as Neonympha areolatus (J. E. Smith, 1797) (type locality USA: Georgia, Chatham Co.), Godart's species number 58, he used "Satyre aréolé" as a French name, and " Satyrus areolatus " as a Latin name ( Godart [1824]). Even more striking an example is "Polyommate W-blanc" for " Polyommatus W-album " ( Godart [1824]: 648). However, for the majority of species, the second word in the French name matched the species epithet more closely than in these cases. Because of that, the situation with two sets of names may be confusing to a casual reader, but Godart meticulously followed this distinction. Only French names were used in the section of the work giving brief diagnoses (e.g., pp. 460–476) ( Godart [1824]). A French name was also used as the title for each species description (e.g., pp. 477–552), immediately following the species number. Latin names were given in the second line of these expanded descriptions, after the French names. Given this distinction between French and Latin names employed by Godart, a deviant spelling "cautheus" [3 rd letter "u", not "n"] occurred only once and in the French name ( Godart [1824]: 465), not in the Latin name, and therefore is irrelevant to zoological nomenclature. The correct (and only) spelling of this species epithet in Godart's work is " cantheus ".

Furthermore, because there was no Fabrician name canthus , there could be no replacement name for it. Therefore, Godart did not propose a "replacement name", he proposed a new name: Satyrus cantheus , and this taxon must have had type specimen(s). Because Godart's description of S. cantheus ends with "(Translation of Fabricius)", it is possible that Godart did not inspect any specimens at all, as also suggested by Cardé et al. (1970): "type ... probably never existed", but based his new species entirely on the Fabrician texts. This hypothesis may not be entirely true, because Godart elaborated on the Fabrician description by mentioning the size of his S. cantheus (species number 56): "Its size is about the same as the previous one", which was species number 55 P. canthus Linnaeus, 1767 (a junior objective synonym of L. eurydice ) ( Godart [1824]). No reference to this size was given in any Fabrician works we studied, and it should have come from some additional knowledge Godart had about the type specimen(s). Whether Godart inspected the actual specimens or not, the name-bearing type currently encompasses a series of all specimens used to constitute Godart's description of cantheus , either directly examined by him, or described by Fabricius in those parts of his texts that were cited and translated by Godart as falling under Godart's concept of cantheus . Among these specimens, there should be at least one specimen possessing, as per its description, a combination of the following four characters: (1) size approximately as L. eurydice ; (2) 3 small indistinct ventral forewing eyespots; (3) the 5 th ventral hindwing eyespot (counting from the apex) being the largest of the six present, and (4) the 6 th spot (by the anal angle) being the smallest. To define the taxon S. cantheus , we hereby designate the syntype that has all 4 characters stated above and was found first (with the earliest date and time) in the search for syntypes of Satyrus cantheus as the lectotype of Satyrus cantheus Godart, [1824] . This designation satisfies the ICZN Code ( ICZN 1999). Our lectotype fulfills the definition of the term as a single specimen selected out of syntypes subsequently to the establishment of the name (ICZN Code Glossary). The designation is individual: it is made for cantheus only (Art. 74.3.). We employ the term " lectotype " (Art. 74.7.1.). Our designation contains information sufficient to recognize this single specimen by the characters given as being the first syntype found in the search for syntypes that possesses all these characters (Art. 74.7.2.). The reason for the lectotype designation is to objectively define this taxon by selecting the specimen that actually agrees with the key characters given in the original description of S. cantheus , and thus represents this species as intended by Godart, promoting stability of nomenclature.

Next, we searched for the lectotype in collections and databases where the types of Fabricius and Godart could be. In particular, N.V.G. inspected the holdings of the Muséum National d'Histoire Naturelle, Paris, France (MNHP), which are known to have a number of Godart types, some with a characteristic "god." signature on their labels; and looked through the relevant parts of the collections of the Natural History Museum, London, UK (BMNH) and the Museum für Naturkunde, Berlin, Germany (ZMHB). We also extensively searched online databases, including the Natural History Museum of Denmark, University of Copenhagen, Denmark (2021), the Hunterian collections, University of Glasgow, Scotland, UK (2021) and the National Museums of Scotland, Edinburgh, UK (2021). Our searches failed to find the lectotype that also has not been found by previous workers, and therefore we believe it became lost. Hence, we proceed herein with a neotype designation. There is an exceptional need to designate the neotype of S. cantheus . Although this name has not been used as valid for over a century and some considered it a misspelling ( Miller and Brown 1981), others a nomen dubium ( Cardé et al. 1970), Viloria (2021) resurrected it and applied it in a manner inconsistent with the original description. To proceed with a meaningful integrative taxonomic revision of Hermeuptychia , it is essential to stabilize the identity of S. cantheus by a neotype that while being in a reasonable agreement with the original description, minimally disrupts currently used taxonomic arrangement and allows more detailed genomic studies of the neotype. DNA work plays a highly significant role in Hermeuptychia studies ( Cong and Grishin 2014; Seraphim et al. 2014; Tan et al. 2021). Therefore, the neotype should facilitate DNA-based studies.

First, we agree with Viloria (2021) that S. cantheus is most likely a Hermeuptychia . Its description is rather similar to the descriptions of other Hermeuptychia species by these authors. Second, in agreement with the original type locality ("North America", which nevertheless could have been an error and simply carried by default from P. canthus , which is L. eurydice ), we aim to select a neotype from North America. North American Hermeuptychia are typically decisively smaller than L. eurydice , and the largest populations are known from the southeastern USA, which is also a general area that contributed type specimens for many early butterfly names. Therefore, we looked for a southeastern USA Hermeuptychia specimen that matches best the original description of S. cantheus . The specimen that we hereby designate as the neotype of Satyrus cantheus Godart, [1824] is the male pictured in Figs. 31, 32 . It largely agrees with the original description: immaculate brown on dorsal side; ventral side paler, with 2 brownish lines on both wings, 3 not very large eyespots on the forewing, 6 on the hindwing, out of which the 5 th is large and the 6 th by the anal angle is smaller. The discrepancies with the original description are: the 5 th eyespot is the same size as the 2 nd eyespot (not larger); the 6 th eyespot, while being small, is larger than the 1 st (not the smallest); and this being large for a USA Hermeuptychia specimen (FW length 17 mm) is still smaller than a typical L. eurydice . However, due to significant variation in Hermeuptychia eyespots, we were satisfied with the reasonable match to the description, and did not invest additional effort into looking for a perfect match. Additional advantage of this specimen is the availability of DNA sequences.

Our neotype of S. cantheus satisfies all the requirements set forth by the ICZN Article 75.3. Namely: 75.3.1. It is designated to clarify the taxonomic identity of Satyrus cantheus Godart, [1824] , which remained undefined until now, impeding research on Hermeuptychia ; 75.3.2. The characters that differentiate this taxon have been given in its original description by Godart ([1824]: 465, 493–494), i.e., wings brown on dorsal side, ventral side paler, with two ferruginous lines spanning both wings, forewing with three weakly defined eyespots and hindwing with six; 75.3.3. The neotype specimen is a male that bears the following six rectangular labels: [St. Petersburg, Fla. | XI-3-1938], [H. E. Wilford | collector], [96], [Ernest Shoemaker | Collection 1956], [DNA sample ID: | 11-BOA-15609E04 | c/o Nick V. Grishin], and [NEOTYPE ♂ | Satyrus cantheus | Godart, [1824] | designated by Grishin] and its partial COI barcode sequence has been deposited in GenBank with accession KJ025563 ( Cong and Grishin 2014); 75.3.4. Our search for the lectotype is described above, but it was unsuccessful, leading us to believe that the lectotype is no longer extant; 75.3.5. The neotype is consistent with what is known about this taxon, in particular with the original description: e.g. ventrally, it has 3 less prominent eyespots on the forewings and 6 eyespots on the hindwings, the 5 th of which is large and the one by the anal angle is small; 75.3.6. The neotype is from USA: Florida: Pinellas Co., St. Petersburg, and the lectotype was stated to be from North America with no other details given about the locality; 75.3.7. The neotype is in the collection of the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA (USNM).

The neotype of S. cantheus is the species known today as Hermeuptychia sosybius ( Fabricius, 1793) ( Cong and Grishin 2014) : it has patches of darker androconia on the dorsal surface of the wings and its COI barcode sequence is identical to that of the H. sosybius neotype. Therefore, Satyrus cantheus Godart, [1824] is a junior subjective synonym of Hermeuptychia sosybius ( Fabricius, 1793) .

Figures

Figs. 19–32. Hermeuptychia specimens. H. canthe: 19–20. neotype ♂, designated herein, Suriname: Gelderland, Suriname River, genitalia slide W. D. F. #306 (Fig. 34), DNA sample NVG-14061C10 [USNM]; 21–22. ♂, Peru: Madre de Dios, Amazonia Lodge, Atalaya, 491 m, 25-Sep-2011, leg. Steve Kinyon, #1907, DNA sample NVG-2649, genitalia NVG140628-17 (Fig. 35i) [USNM]; 23–24. ♀, Guyana: Region 9, Kanuku Mts., Nappi Creek, 03°20.7'N 59°34.2'W, 500'-1000', 21-Feb-10-Mar-1999, leg. S. Fratello, R. Hanner, S. Hendricks & R. Williams, USNM ENT 00232265, DNA sample NVG-2675, genitalia NVG140628-43 (Fig. 35l–m) [USNM]; 25–26. ♂, Colombia: Caqueta, Florencia, 6-Aug-1976, fairly wet hillside forest, leg. J. A. Scott, DNA voucher NVG-2798; 27–28. ♀, Suriname: Commewijne, De Nieuwe Grond, secondary forest, 2-Jun-1982, leg. Olle Pellmyr, DNA sample NVG-2644, genitalia NVG140628-12 (Fig. 35j–k) [USNM]; 29–30. illustrations of type specimens from the original description, not to scale (Hübner [1811]). 31–32. H. cantheus neotype ♂, designated herein, USA: Florida, St. Petersburg, 3-Nov-1938, leg. H. E. Wilford, DNA sample 11-BOA-15609E04 [USNM].