identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03B787FAA55471704E85FDB3BF7E9DE0.text	03B787FAA55471704E85FDB3BF7E9DE0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Papilio marcus Fabricius 1787	<div><p>Papilio marcus Fabricius, 1787</p> <p>The specific epithet marcus was introduced by Johann Christian Fabricius in his Mantissa insectorvm sistens species nvper detectas adiectis synonymis, observationibvs, descriptionibvs, emendationibvs (Fabricius 1787: 87). Like other butterfly species described in Fabricius (1787), the name appeared in combination with the generic name Papilio Linnaeus, 1758, and this taxon is currently widely regarded as a valid species in the Neotropical skipper genus Troyus Warren &amp; Turland, 2012 (Zhang et al. 2019: 8, Fig. 13; Fig. 1A), although it must be noted that a few early studies did consider this taxon as conspecific with P. phyllus Cramer, 1777 (see below). Fabricius described P. marcus based on an unspecified number of specimens in the collection of “v[on]. Rohr” (i.e., Julius Philipp Benjamin von Rohr [1737-1793]), with an unknown sex from Cayenne [French Guiana]. For over 100 years, this species-group name was associated with the skipper genus Vettius Godman, 1901 (Godman 1901: 589), and the combination Vettius marcus was used in many important publications regarding skipper classification (e.g., Evans 1955: 182; Mielke 2004: 84; 2005: 1350). In particular, a number of publications associated this name with illustrations of the adults starting with Geyer (1832: figs 725, 726; Fig. 2), who placed marcus under the generic name Cobalus Hu ̈bner, [1819] (Geyer 1832: 30; Fig. 2). Subsequently, the following publications illustrated the wing pattern: Draudt (1923: fig. 188g, ventral surface); Hayward (1934: fig. 21, ventral surface; 1950: pl. 24, figs 14, 15, both surfaces); Lewis (1973: figs 24, 25; 1975: figs 24, 25); Maza (1987: pl. 67, fig. 5, dorsal surface); Canals (2003: figs, both surfaces); Brown &amp; Freitas (2000: fig., ventral surface); Glassberg (2007: 245 fig., live picture, ventral surface); Cock (2009: figs 7-9, both surfaces, live picture); Garwood et al. (2009: 328, fig., live picture, ventral surface); Garwood &amp; Lehman (2013: 224, fig. [identical individual illustrated in preceding article]) and/or its male genitalia: Godman (1900: pl. 102, fig. 10); Hayward (1934: fig. 19; 1950: pl. 12, fig. 14); Evans (1955: pl. 66). The illustrations of genitalia in these studies reflect individuals with characters currently corresponding to the concept of “ marcus ” in the literature (Fig. 1B). In addition, both the wing illustrations and genitalia are consistent with each other, thus revealing a long-standing consensus about the identity of the skipper taxon represented by the name “ marcus ”.</p> <p>Papilio marcus Fabricius, 1787 is in reality a junior primary homonym of Papilio marcus Schaller, 1785, a name currently validly employed for a species of the popular butterfly genus Morpho Fabricius, 1807 (Nymphalidae: Satyrinae) (Lamas 2004). Because these names apply to taxa not considered congeneric after 1899, the case needs to be referred to the International Commission on Zoological Nomenclature for a ruling under the plenary power regarding this homonymy (ICZN 1999:Article 23.9.5). Notwithstanding this situation, an examination of the syntype of Papilio marcus Fabricius, 1787 currently housed at ZMUC (see also discussion below; Fig. 1C) raised some questions as to its identity. Contrary to the current concept of “ marcus ”, as exemplified in the above references, the ventral surface of this male syntype in ZMUC is lacking yellow scaling at the distal side of hindwing M 3 -Cu 1 and Cu 1 -Cu 2, as well as not having whitish markings in the basal side of hindwing cells Cu 2 -2A and 2A-3A. The white marking in the ventral hindwing discal cell does not reach the distal (outer) margin in the figured specimens, whereas this corresponding marking in the syntype of P. marcus reaches the distal margin of the hindwing and, moreover, appears yellowish. Another distinctive difference is the presence of fulvous scaling along the costal and inner margins of the ventral hindwing in the syntype in Copenhagen, whereas this reddish marking is not visible on the ventral hindwing in individuals we perceive as Troyus “ marcus ”.</p> <p>In fact, as indicated by one of the labels associated with this syntype in Copenhagen, these wing pattern characters suggest its identity to match that of the species currently known as Vettius phyllus (Cramer, 1777). Papilio phyllus was described by Pieter Cramer based on an unspecified number of specimens of an unknown sex from Suriname, owned by Enricus Fredericus Alberti (Cramer 1777: 122). Although the number of specimen(s) is not explicitly mentioned, Cramer’s original description refers to the specimen as a singular (i.e., “ this little Plebejd.. ”), thus he likely relied on a single specimen. The illustrations of this taxon on pl. CLXXVI [176], accompanying the original description, depict the dorsal surface on Fig. B and the ventral surface on Fig. C. Based on the examination of the original illustrations by Cramer housed at NHMUK (these drawings were used to produce the hand-colored illustrations for the published book; see Chainey 2005), the apparent lack of three whitish, or silver, markings on the ventral hindwing cells M 2 -M 3, M 3 -Cu 1, and Cu 1 -Cu 2 is not in accordance with V. phyllus; however, this illustration of the ventral surface is largely in agreement with the syntype housed at ZMUC and distinguishable from the aforementioned illustrations of “ marcus ” by the suite of characters mentioned above. To our knowledge, the syntypes of P. phyllus have not been located at RMNH (e.g., Gernaat et al. 2012; B. Hermier, pers. comm.), where some of Cramer and Stoll’s types can be found. The Rothschild collection housed at NHMUK also includes types by Cramer, mostly from the van Lennep collection. These were subsequently purchased by Cajetan Felder, although we were unable to find them in London, and thus we are unable to extend the discussion regarding application of this species-group name beyond the original description by Cramer. Based solely on a phenetic comparison, we are able to exclude a number of available names potentially applicable to the syntype of phyllus, and settle the nomenclature accordingly.</p> <p>Among the taxa discussed here, the fulvous costal and inner margins of the hindwing, discernible in the marcus syntype in Copenhagen, is a feature observed in what are currently perceived as Vettius phyllus and two closely related species, namely V. chagres Nicolay, 1973 and V. mitsuko Nakahara, Nakamura &amp; MacDonald, 2020, and these three taxa form a clade (Nakahara et al. 2020). However, V. chagres and V. mitsuko are both known only from Central America and are externally distinguishable from V. phyllus (see Nakahara et al. 2020) and thus are unlikely to represent the syntype of P. marcus, which is supposedly from Suriname. As evidenced by Zhang et al. (2019), seven species currently placed in Troyus - T. turneri Warren &amp; Turland, 2012, T. fantasos (Cramer, 1780), T. onaca (Evans, 1955), T. aurelius (Plötz, 1882), T. “ marcus ” (Fabricius, 1787), T. diversa (Herrich-Schäffer, 1869), and T. drova (Evans, 1955) - are recovered as a clade, sister to Monca Evans, 1955 and phylogenetically distantly related to Vettius. Despite some of these taxa overall resembling V. phyllus, perhaps explaining why they were previously considered congeneric under the genus Vettius (e.g., Mielke 2004, 2005), none of these species, except for Troyus “ marcus ” (see below for further discussion), possess a fulvous costal and/or inner margin of the hindwing. It therefore seems reasonable to identify the syntype of P. marcus Fabricius, 1787 as P. phyllus based on available species-group names.</p> <p>The identity of the aforementioned syntype in Copenhagen as P. phyllus is also supported by the original description by Fabricius: Turton’s (1806: 169) translation - [Papilio Hesperiae Urbicolae] Marcus. Wing without tails brown with hyaline spots; lower-ones beneath with 2 yellowish fillets, the thinner margin fulvous. Inhabits Cayenne; small. Antennae hooked, Lower-wings with a white thinner margin and large spot in the middle above ”. Papilio phyllus, as perceived by many authors throughout history (e.g., Cramer 1777: pl. [176], figs B, C; Hayward 1934: pl. 11, figs 19, 20; Garwood et al. 2009: 328, fig.), is a widespread species in the Neotropical region and this taxon is the type species of Vettius by original designation of Godman (1901).</p> <p>Latreille [1824: 753] realized that Papilio phyllus and P. marcus Fabricius, 1787 were conspecific based on the fact that the specimen he had in his hands to prepare the description of P. marcus Fabricius, 1787 possessed the bluish dorsal spots and fulvous inner hindwing margin found in P. phyllus. These two features mentioned by Latreille are present in P. phyllus and not T. “ marcus ”, as partly discussed above. The latter feature, mentioned in the description of P. marcus by Fabricius, likely influenced Latreille’s decision to regard these taxa as conspecific, although he considered P. marcus Fabricius, 1787 to be given priority for an unknown reason. Subsequently, several authors considered Papilio phyllus and P. marcus Fabricius, 1787 to be conspecific, starting with Westwood (1852: 527; P. marcus Fabricius, 1787 again as the name taking precedence), followed by Butler ([1870: 273]; P. phyllus as a senior name), Kirby (1871: 590; P. phyllus as a senior name). Plötz (1882: 454) recognized the synonymy of Latreille’s ([1824]) “ marcus ” and P. phyllus, although he regarded P. marcus Fabricius, 1787 (= phyllus sensu Kirby) as a discrete taxon. Godman (1900: 585) also noticed these synonymous treatments of these two taxa, yet he stated that this was an erroneous taxonomic proposal. The synonymy by these authors is also reflected in Mielke (2005), but it is unfortunate that this fact did not influence a number of other authors who considered P. “ marcus ” to be a discrete taxon.</p> <p>Zimsen (1964: 546) listed a single specimen for the type material of P. marcus Fabricius, 1787 as “in Kiel” (i.e., ZMUK). Fabricius’ personal collection belonging to the ZMUK was on a long-term loan to the ZMUC from 1958 to 2013, where the Sehested-Tønder Lund collection has been housed since the early 1800s, and they were kept separate (Tuxen 1967; O. Karsholt, pers. comm.). Subsequently, it was negotiated that all specimens of Coleoptera should be transferred to ZMUK, whereas the rest of the collection should be kept in Copenhagen. As a result, all type material of Lepidoptera from Fabricius’ personal collection and the Sehested-Tønder Lund collection are now property of the ZMUC and housed accordingly.</p> <p>Julius von Rohr, a German botanist in the Dutch colonial service and residing in Saint Croix, studied cotton culture and embarked on a journey to the West Indies and the northern coast of South America, including Cayenne (i.e., the type locality of P. marcus Fabricius, 1787) (Millspaugh 1902). Hopkins (1999) also noted that von Rohr lived in the American tropics for 30 years, supporting the fact that he was able to directly obtain or collect specimens that were later used by Fabricius to describe P. marcus. According to Papavero (1971), von Rohr was asked by the government to travel to Jamaica, Puerto Rico, Lesser Antilles, as well as along the coast of the Guianas, and later sent back a large insect collection to Copenhagen. Existing literature also supports the claim that Julius von Rohr sent herbarium samples to Niels Tønder Lund (Hopkins 1999), and, directly or indirectly, to Martin Vahl in Copenhagen (Fryxell 1969; Millspaugh 1902; Hopkins 2016), as well as to Joseph Banks in London (Hopkins 2016). Indeed, Fryxell (1969) studied the cotton plant genus Gossypium L. (Malvaceae) based on von Rohr’s materials in Copenhagen, which were received by Martin Vahl and remained there until his study. Hence, if insect specimens were shipped with plant materials, it is reasonable to assume that they should have ended up in Copenhagen and entered the Sehested-Tønder Lund collection, unless Banks was the recipient of von Rohr’s insects. Indeed, this taxon did not appear in Aurivillius’ (1898) list of Fabrician Lepidoptera in Copenhagen. Zimsen (1964) lists 267 insect species described by Fabricius based on materials in von Rohr’s possession, although only 13 species appear to be housed in Copenhagen, with Kiel mentioned as the repository for three of them. Henriksen (1921) described the donation of two drawers to the Danish Natural History Society by Sehested and Tønder Lund, which partly included insect materials from von Rohr, and these may have been reflected as “in Copenhagen” in Zimsen (1964). We have reviewed the correspondence between von Rohr and Joseph Banks (Dawson 1958), and consulted with a knowledgeable historian, Daniel Hopkins (University of Missouri - Kansas City, USA) (pers. comm.), who took notes from official records of various Danish government agencies related to colonial, commercial, and financial affairs. These are all the records of correspondence von Rohr had with relevant people in Europe, in its original handwritten form, in the Danish National Archives, and we used these to trace von Rohr’s movements and to investigate how the specimen might have ended up in Kiel with Fabricius. Relevant passages include the following.</p> <p>August 30, 1784: von Rohr wrote to Joseph Banks from Cayenne, where he was living for a year or so.</p> <p>February 11, 1785: von Rohr is reported to still be in Cayenne, asking the Danish government for funding to return home.</p> <p>When von Rohr finally gets back to St. Croix, he learned that the clove seedlings he forwarded from Cayenne were destroyed by a hurricane [in August 1785] and he turns his attention to repairs of official buildings in St. Croix.</p> <p>May 31, 1786: a communication from von Rohr in St. Croix.</p> <p>December 22, 1786: von Rohr receives a prize for his work in St. Croix.</p> <p>October 29, 1787: the colonial office in Copenhagen refers to von Rohr, in St. Croix, where he has been given another year off from his regular duties to work on a book about cotton cultivation.</p> <p>In addition, there are official communications from von Rohr in St. Croix on April 28, 1787 and July 11, 1787.</p> <p>Fabricius moved to Kiel from Copenhagen in 1775 and remained there until his death in 1808 (Tuxen 1967). Although evidence exists that Fabricius visited Copenhagen periodically starting from 1796, it is unclear whether he visited Copenhagen prior to 1787 to acquire specimen(s) he used to describe P. marcus. Nevertheless, considering the proximity of Kiel and Copenhagen, the possibility of Fabricius obtaining specimens in Copenhagen, either directly or indirectly, and ending up in Kiel is likely high. On the other hand, considering the records described here, it seems unlikely that von Rohr returned to Copenhagen, or elsewhere in Europe, in the period in question, taking into account the slowness of travel at that time. As stated in Papavero (1971), he most likely sent insects to Europe, including P. marcus, perhaps from Cayenne, in 1784 or 1785. Considering von Rohr’s correspondence with Banks during his time in the Antilles and the northern coast of South America, he might have sent specimens of P. marcus to Banks, which eventually ended up with Fabricius. Other possible recipients of von Rohr’s insect specimens include Martin Vahl, Tønder Lund and Sehested, both at Copenhagen.</p> <p>Hopkins (2016) stated “ Von Rohr was travelling with three local Amerindian guides and two black slaves of his own, he wrote; one of his slaves specialised in collecting insects, the other in plants ”. An examination of correspondence between Benjamin Smith Barton and von Rohr, which took place in Philadelphia, USA, in 1793, when von Rohr visited a number of scientists, suggests that these enslaved collectors who travelled with von Rohr were named Julius and Marcus. If Marcus collected a specimen, or specimens, which Fabricius later used to describe P. marcus. it is possible that the specific epithet is based on von Rohr’s enslaved insect collector. Nevertheless, this is no more than speculation.</p> <p>We have consulted with Ole Karsholt (ZMUC) to confirm that there is only a single specimen of P. marcus in Fabricius’ personal collection housed in Copenhagen, and thus the number of syntypes is in agreement with Zimsen’s (1964) type catalogue. The original description by Fabricius does not contradict this unique specimen of Papilio marcus in Copenhagen, and most likely was considered by Fabricius to represent P. marcus. Thus, it is suitable to serve as the objective standard for this named entity. Since providing evidence for the existence of additional syntype (s) is not possible, we therefore accept this male specimen as an authentic syntype of P. marcus Fabricius, 1787. Consequently, we designate this specimen as the lectotype of P. marcus Fabricius, 1787 in order to settle the nomenclature and identity, as perceived in the present study (i.e., as a junior synonym of P. phyllus) (lectotype designation). Although three subspecific names have been associated with P. phyllus (Mielke 2005), the marcus lectotype from Cayenne is likely to represent the nominate race, considering that the type locality of P. phyllus lies in its neighboring Suriname.</p> <p>Given that we identified the syntype of P. marcus Fabricius, 1787 as P. phyllus, we review species-group names historically associated with P. “ marcus ” in order to find a name that suits the current concept of P. “ marcus ”, as outlined above.</p> </div>	https://treatment.plazi.org/id/03B787FAA55471704E85FDB3BF7E9DE0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Nakahara, Shinichi;Zilli, Alberto;Calhoun, John V.;Espeland, Marianne;Padrón, Pablo Sebastián;Grishin, Nick V.	Nakahara, Shinichi, Zilli, Alberto, Calhoun, John V., Espeland, Marianne, Padrón, Pablo Sebastián, Grishin, Nick V. (2022): Resolving two centuries of mistaken identity: Reinterpretation of Papilio marcus Fabricius, 1787 (Lepidoptera: Nymphalidae, Hesperiidae). Zootaxa 5195 (3): 241-255, DOI: https://doi.org/10.11646/zootaxa.5195.3.3
03B787FAA551717E4E85FBF5BFE69C48.text	03B787FAA551717E4E85FBF5BFE69C48.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hesperia aurelius Plotz 1882	<div><p>Hesperia aurelius Plötz, 1882</p> <p>Plötz (1882: 455) described Hesperia aurelius based on an unspecified number of specimens from Brasilien (= Brazil). Despite the fact that the original description in German was not accompanied by illustrations, the existence of drawings is indicated in the original description (as t. 444), as well as evidenced from an external source (Godman 1907). Plötz’s rather detailed description in the form of a dichotomous key (see below) suggests that his H. aurelius is in accordance with the current concept of Troyus aurelius (e.g., Godman 1900: pl. 102, figs 11, 12; Draudt 1923: pl. 188h; Zhang et al. 2019: fig. 13), largely by mentioning the snow-white tornus and anal margin of the ventral hindwing. As discussed below, the whereabouts of the type material and illustrations for the skipper species discussed in Plötz (1882) are currently unknown (but see below). Types of some African butterfly taxa described in Plötz (1880) are documented in Libert (2004) and are currently deposited at the ZIMG. Lara Lopardo (pers. comm.), the present collection manager at the ZIMG, examined Plötz’s collection of 157 skipper specimens housed in Greifswald, including 30 type specimens, and confirmed that the type material for H. aurelius is not deposited at Greifswald.</p> <p>The name aurelius subsequently appeared in combination with the generic name Carystus Hübner, [1819] in Godman (1900: 585), where both surfaces of a female specimen were figured (pl. 102, figs 11, 12). The rationale of associating the species-group name aurelius to the illustration by Frederick Godman remains unknown. At some point between 1882 and 1907, Godman did receive Carl Plötz’s drawings (Godman 1907), but little evidence exists to determine when he examined them. Actually, in the NHMUK archives there exists a copy of Plötz’s numbered drawings that Godman commissioned to Horace Knight and other artists to keep pictorial evidence of the taxa illustrated by the German entomologist. However, in the collation there are many leaps with regard to Plötz’s sequence, as Godman evidently requested copies only of the taxa lesser known to him. Unfortunately, neither aurelius nor lyrcea Plötz, 1882 (see below) were copied, likely because Godman had no doubts about their identity.</p> <p>The collation is tentatively dated as from 1912 and ought to cover only the American taxa, as per the title on its binding (“ Copies of Plötz’s unpublished figures of American Hesperiidae ”), but in reality, a number of species from other zoogeographical regions are also illustrated. Godman’s (1907) statement “ I have recently had an opportunity of examining …. by the late Carl Plötz ”, suggests he was only able to examine these plates just prior to Godman (1907), and not during the course of preparing Godman (1900). Having died in 1886, Carl Plötz was not able to provide his illustrations to Godman in this time-frame, and we are unable to trace how drawings ended up with Godman. Plötz’s originals seem to have ended up in ZSM, where a number of them are pinned inside the Hesperiidae drawers. It is therefore likely that additional drawings may be in the massive and yet unsorted miscellaneous collection of Plötz’s documents, letters, manuscripts etc. in the ZSM library (A. Hausmann &amp; U. Buchsbaum, pers. comm.).</p> <p>Godman’s (1900) illustration of aurelius on pl. 102, figs 11, 12, appears to be in accordance with how the species is recognized today. largely by having a white posterior band on the hindwing covering most of the inner margin (in accordance with Plötz’s (1882) description). Despite being treated as a valid species, Godman (1900) noted that “ Carystus ” aurelius is merely a form of C. marcus since he could detect no difference in the male genitalia of these two taxa. This species-group name has since been considered subspecific by some authors (e.g., Evans 1955; Mielke 2004, 2005) and species-level by others (e.g., Draudt 1923; Steinhauser 1975), although none of these authors provided meaningful arguments towards its taxonomic status. Evans (1955) did not illustrate the genitalia for aurelius, perhaps for the same reason Godman (1900) did not figure them, namely that they are identical to P. “ marcus ”. The series of T. aurelius at NHMUK, 46 of which were examined by Evans (1955), appear to consist of specimens that fit the current concept of Troyus aurelius. These specimens, as well as literature references where the specific epithet “ aurelius ” was associated with figures (see above), show a consensus in the characters throughout time regarding the identity of the skipper taxon represented by the name “ aurelius ”. Furthermore, Zhang et al.’s (2019) genetic data supports its species-level status, rather than as a subspecies of Troyus “ marcus ”. Finally, the ambiguous application of the species-group name justifies our consideration of the specific epithet “ aurelius ” as “in use” and should not be applied to the current concept of P. “ marcus ”, compared to the name “ phyllides ” discussed below.</p> <p>Descending order of the relevant key couples and taxa in Plötz’ (1882) key are as such:</p> <p>A, B, C, etc. a., b., c., etc. ○, ○○, ○○○, etc. ‒, ‒ ‒, ‒ ‒ ‒, etc. ^, ^^, ^^^, etc. |, | |, | | |, etc. ˅, ˅˅, ˅˅˅, etc. §, §§, §§§, etc.</p> <p>A. Antennae almost always more than half the length of the forewing</p> <p>b. [p. 323] Wings with hyaline spots. [opposed to: a. Wings without hyaline spots.</p> <p>○○ [p. 324] The palpi are not red. [opposed to: ○ Upperside black, The body and the base of the wing roots with glossy blue or green long scales (hairs), forewing with white glass spots. Palpi red.]</p> <p>‒ ‒... ‒ [p. 453] Upperside black. FW with a pale spot in cell 1, a white hyaline spot in cell 2, one in cell 3, a half or a point in the middle cell and two points in cells 6 and 7. HW with a large white or yellow spot in the middle and against the anal margin such a ray. Beneath have the FW a large pale yellow spot in front of the apex or at least in cells 4 and 5, the HW are yellowish, vein 6 is mostly very darkened in its entire length and the anal angle is darkened. [opposed to all the other ‒ ‒... ‒ couples of the key, with diverse comments.]</p> <p>^^ [p. 454] Forewing with a dot or line on upper part of the discoidal cell. Hindwing underside with a narrow brown longitudinal stripe. [opposed to: ^ FW with half a spot on the inner margin of the central cell and a blue stripe on the rear edge of the wings. HW above with a large white central spot and two blue rays, beneath with a broad brown longitudinal stripe along vein 6, brownish anal margin and tornus. Collar red.]</p> <p>| [p. 454] Discal spot on the hindwing upperside sharply limited [yes], tornus blackened on the underside. [opposed to: || Hindwing above with a white large central spot that flows into the rear corner up to vein 2, below with the tornus and the anal margin snow-white. Forewing below with a large pale yellow spot at the apex. Collar red.]</p> <p>˅ [p. 454] All the spots are white on the upperside. [opposed to: ˅˅ The forewing spot in cell 1, the spot and the streak on the hindwing, the fringes, and, on the underside, the costa of the forewing are pale sulfur yellow. Collar black.]</p> <p>§ [p. 454] The pale spot beneath at the apex of the FW is washed out [=fading]; on the HW the white middle spots elongated. → Marcus [opposed to: §§ On the forewing underside, the hyaline spots in cells 4 and 5 are sharply delimited; on the hindwing, the white center spot is rounded on the rear flights. Collar red. Club of the antenna with a white point.]</p> <p>§§ [p. 454] The pale spot beneath in cells 4 and 5 of the FW is sharp delimited; on the HW the white spot is rounded. Collar red. Head dotted with white. → Lyrcea [opposed to: § On the forewing underside, the lower hyaline apical spot is washed out; on the hindwing, the white center spot is stretched.]</p> <p>|| [p. 455] HW dorsal with a white large central spot that has “flowed out” into the tornus angle up to vein 2, beneath at the tornus and anal margin snow-white. FW beneath with a large pale yellow spot in front of the apex. Neck collar red.→ Aurelius [opposed to: | Discal spot on the hindwing upperside sharply limited [yes], tornus blackened on the underside.]</p></div> 	https://treatment.plazi.org/id/03B787FAA551717E4E85FBF5BFE69C48	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Nakahara, Shinichi;Zilli, Alberto;Calhoun, John V.;Espeland, Marianne;Padrón, Pablo Sebastián;Grishin, Nick V.	Nakahara, Shinichi, Zilli, Alberto, Calhoun, John V., Espeland, Marianne, Padrón, Pablo Sebastián, Grishin, Nick V. (2022): Resolving two centuries of mistaken identity: Reinterpretation of Papilio marcus Fabricius, 1787 (Lepidoptera: Nymphalidae, Hesperiidae). Zootaxa 5195 (3): 241-255, DOI: https://doi.org/10.11646/zootaxa.5195.3.3
03B787FAA55F717D4E85FD56BCFF9F34.text	03B787FAA55F717D4E85FD56BCFF9F34.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hesperia lyrcea sensu Plotz 1882	<div><p>Hesperia lyrcea Plötz, 1882</p> <p>Carl Plötz described Hesperia lyrcea simultaneously in Entomologische Zeitung (Stettin) in a page preceding his description of H. aurelius, based on an unspecified number of specimens that were also from Brasilien (= Brazil) (Plötz 1882: 454). After examining illustrations prepared by Carl Plötz (see above under aurelius), Godman (1907) synonymized H. lyrcea under P. “ marcus ”. Hesperia aurelius, which supposedly appeared in Carl Plötz’s plates for his new skipper species, was not commented on in Godman (1907), perhaps because Godman (1900) already discussed its conspecificity with P. “ marcus ” as discussed above. Draudt (1923: 979) followed this synonymy by Godman. Evans (1955: 183) considered lyrcea as a subspecies of Vettius diversa (Herrich-Schäffer, 1869), a taxon initially described as Cobalus diversa. Evans (1955) refers to figures for “ diversa ” (i.e., diversus) in Godman &amp; Salvin (= Godman 1900) and Seitz (= Draudt 1923) as lyrcea without any explanation as to these associations. This decision by Evans (1955) was presumably based mainly on the fact that lyrcea possesses dark veins on the hindwing (thus dividing yellowish ventral hindwing markings), whereas veins are not dark in P. “ marcus ”, a character used to distinguish V. diversa from V. marcus in Evans (1955). The internal view of the valva figured for lyrcea (Evans 1955: pl. 66) illustrates differences in the harpe and overall shape of the valva compared to other potential congeners figured on the same plate, including V. marcus and other subspecies of V. diversa.</p> <p>A review of specimens at NHMUK reveals 32 specimens of lyrcea that show consistent differences between Troyus “ marcus ” mainly by Rs, M 3 and Cu 1 of the ventral hindwing being dark and thus dividing yellowish or whitish markings. Evans (1955) examined 34 specimens in total, 33 from Ecuador and a single male from Peru. There exists space for two specimens in the corresponding section of the drawer in the NHMUK, thus it is reasonable to consider that these were the specimens examined by Evans to investigate the identity of lyrcea. Furthermore, we found four specimens without abdomens, as well as two specimens with apparently intact abdomens with their genitalia glued on a card that is pinned beneath them. They were presumably dissected by humidifying the abdomens and extracting the genitalia without removing the abdomens from the specimens. In the preceding work (Evans 1949), Norman D. Riley stated that Evans’ intention was to provide “diagrammatic” illustrations of male genitalia, and given that any of these six individuals (summary of multiple, or none of them) might have been used to establish the identity of lyrcea sensu Evans (1955), it is impossible to draw conclusions as to how Evans (1955) identified this taxon. Carl Plötz’s key leading to lyrcea is questionable as to its reliability in distinguishing this taxon from congeners, namely by stating “ the pale spot beneath in cells 4 [M 2 -M 3] and 5 [M 1 -M 2] of the FW is sharp delimited [opposed to “ the lower hyaline apical spot is washed out”] ”. Even based solely on specimens in the drawer in the NHMUK, these spots in the ventral hindwing cells M 2 -M 3 and M 1 -M 2 do not appear to show inter-specific differences in these taxa. Plötz therefore likely examined limited numbers of specimens, or perhaps single specimens, to prepare his descriptions. Despite the lack of type material and original illustrations, the most up-to-date comprehensive catalogue for Hesperiidae retains the taxonomic status of lyrcea as a subspecies of Vettius diversa (Mielke 2004, 2005), in accordance with Evans (1955). As stated above, our attempt to locate syntype (s) at ZIMG and MFNB did not yield any specimens which could represent Plötz’s type material relevant to the present study. While there appears to be no ambiguity as to Evans’ (1955) concept of lyrcea, we are unable to apply the specific epithet lyrcea sensu Plötz (1882) mainly due to the lack of type material. In the present study, we refrain from drawing any conclusions regarding the identity of Hesperia lyrcea, which may or may not represent a species whose concept matches P. “ marcus ”. Under the current circumstances, we will not designate a neotype for H. lyrcea, and consequently do not regard this taxon as conspecific with P. “ marcus ”.</p> </div>	https://treatment.plazi.org/id/03B787FAA55F717D4E85FD56BCFF9F34	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Nakahara, Shinichi;Zilli, Alberto;Calhoun, John V.;Espeland, Marianne;Padrón, Pablo Sebastián;Grishin, Nick V.	Nakahara, Shinichi, Zilli, Alberto, Calhoun, John V., Espeland, Marianne, Padrón, Pablo Sebastián, Grishin, Nick V. (2022): Resolving two centuries of mistaken identity: Reinterpretation of Papilio marcus Fabricius, 1787 (Lepidoptera: Nymphalidae, Hesperiidae). Zootaxa 5195 (3): 241-255, DOI: https://doi.org/10.11646/zootaxa.5195.3.3
03B787FAA55C717C4E85FE8ABAC79D3C.text	03B787FAA55C717C4E85FE8ABAC79D3C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Vettius phyllides Rober 1925	<div><p>Vettius phyllides Röber, 1925</p> <p>The species-group name phyllides was established based on a single male and female individuals from Santa Catarina [Brazil] (Röber 1925: 99). The first half of the original description serves as the diagnosis to distinguish Vettius phyllides from V. phyllus based on wing pattern differences: “ I have one pair of this species from Santa Catharina [Brazil], and also one pair of phyllus from the same locality before me, both species are similar sized. The forewings of phyllades [sic] are somewhat narrower, and the posterior half of the hindwings is more curved in than in phyllus. The hyaline spots are smaller in phyllades [sic] than in phyllus, the phyllides female has two post-apical spots. The hind wings show clean, broad white, middle bands (without blue coloration) that start before the M 3 (=hintere Radialis) and reach until the above the 3A (=Innenrandrippe), but is divided by the black 2A (=Submediana); the fringes on the inner margin and in the anal angle are strikingly white and long; the underside of the forewings is almost identical in both species, but the white subapical drawings are more developed in phyllides, which is also the case for the light spot at the inner margin. The underside of the hindwings is white with yellow tint on the anal part; from the wing root stretching along the M 3 (=Mediana?) and the M 1 (= vordere Radialis), as well as at the end of the discoidal cell, from off the 2A to the outer margin there is one narrow grey band, respectively, which both unite in a large, proximally not clearly differentiated spot, at the outer margin between M 1 and Cu 1; in the anal angle, there, however, is a large, dark green distally and proximately undifferentiated spot set apart from the margin. The color of the upper side of the body is identical to that of the wings, but with white hairs, the collar brown, a large white spot in front of each antenna, palpi white with bluish scales; underside of the thorax and body white, and so are the legs; antennae as the wings, the clubs with yellowish underside.”</p> <p>Subsequently, Evans (1955: 182) synonymized V. phyllides under Papilio “ marcus ” without providing justification for this taxonomic rearrangement. He apparently examined specimens from southern Brazil, Paraguay, and Argentina, and they enabled him to reach a conclusion regarding the identity of V. phyllides, given its type locality of Santa Catarina, Brazil. This status as a junior synonym of Papilio marcus Fabricius, 1787 was followed by subsequent authors (e.g., Bridges 1983, 1994). In particular, Mielke (1989: 138) mentioned the examination of a pair of syntypes housed at SMT and designated the male syntype as a lectotype (Fig. 1D), consequently treating the name as a junior subjective synonym of P. marcus Fabricius, 1787. The information regarding the lectotype and paralectotype of V. phyllides, including associated labels, is also reflected in Nekrutenko (2003: 87). There exists no revisionary work on relevant groups to date, and the status of V. phyllides as a junior subjective synonym of Papilio marcus has been retained in all the literature regarding Hesperiidae classification (Mielke, 2004, 2005). The name was therefore never treated as valid subsequent to Evans (1955).</p> <p>Phenotypic inspection and genomic sequencing of the V. phyllides lectotype reveals that it is not conspecific with V. phyllus, and instead belongs to the genus Troyus (Fig. 3, red and magenta). The two genera Vettius and Troyus are not sisters, each being recovered as sister to other genera, Thoon Godman, [1900] and Monca Evans, 1955, respectively, than to each other (Fig. 3). Because P. marcus is a junior subjective synonym of V. phyllus, and V. phyllides is not even congeneric with V. phyllus, V. phyllides cannot remain as a synonym of P. marcus. Reviewing all available names in Troyus, we were not able to find one that could be applied to V. phyllides. Therefore, we propose that Troyus phyllides is a species-level taxon (reinstated status, new combination). This species has been frequently misidentified as Troyus marcus: one such specimen from Suriname is included in the tree (Fig. 3). Finally, the phylogenetic tree reveals that Troyus aurelius is sister to T. phyllides (Fig. 3, red and blue).</p> <p>In conclusion, given the circumstances surrounding the species-group names associated with Papilio marcus Fabricius, 1787, we propose the following three taxonomic changes:</p> <p>(1) Treatment of Papilio marcus Fabricius, 1787 as a junior subjective synonym of P. phyllus Cramer, 1777 (new synonym). By virtue of article 23.9.5 of ICZN (1999), the current placement of Papilio marcus Fabricius, 1787 in Vettius Godman, 1901 was already preventing its automatic replacement as a junior primary homonym of Papilio marcus Schaller, 1785 (now in Morpho Fabricius, 1807), so pending referral of the case to the ICZN, prevailing usage had to be maintained. Removing P. marcus from use as a valid taxon following the above synonymy also eliminates referral to the ICZN as Article 23.9.5 applies only to species-group names “in use”.</p> <p>(2) Removal of the name Vettius phyllides from the synonymy of P. marcus and consideration of this specific epithet as a name to represent a concept regarded by the name Papilio “ marcus ” for nearly 200 years, following Article 23.3.5 of ICZN (1999) (reinstated status). The rationale of applying phyllides to P. “ marcus ” is based on the fact that these two specific epithets have been associated with each other since Evans (1955), and continued to be considered as conspecific in the recent comprehensive catalogue of Hesperiidae (Mielke 2004, 2005). Furthermore, the genomic sequencing data supports the lectotype of V. phyllides as conspecific with specimens that have been misidentified as P. marcus Fabricius, 1787. On the other hand, as explained above, aurelius and other relevant species-group names currently associated with the genus Troyus are no longer associated with Papilio “ marcus ” and, importantly, type material is not available for these taxa to settle their nomenclature. In addition, genetic data exists to support the specific status of these names, which are currently in use (Zhang et al. 2019). We argue that the application of “in use” species-group names to different taxa would cause confusion, and we avoid this situation by proposing a replacement name based on a taxon which has been considered to be a junior subjective synonym. Name-bearing types exist for both Papilio marcus Fabricius, 1787 and Vettius phyllides Röber, 1925, allowing the removal of any ambiguity regarding the application of these names.</p> <p>(3) Proposal of the combination Troyus phyllides (new combination) as a name to refer to a skipper species known as Vettius marcus for over 100 years until its recent taxonomic change in Zhang et al. (2019).</p> </div>	https://treatment.plazi.org/id/03B787FAA55C717C4E85FE8ABAC79D3C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Nakahara, Shinichi;Zilli, Alberto;Calhoun, John V.;Espeland, Marianne;Padrón, Pablo Sebastián;Grishin, Nick V.	Nakahara, Shinichi, Zilli, Alberto, Calhoun, John V., Espeland, Marianne, Padrón, Pablo Sebastián, Grishin, Nick V. (2022): Resolving two centuries of mistaken identity: Reinterpretation of Papilio marcus Fabricius, 1787 (Lepidoptera: Nymphalidae, Hesperiidae). Zootaxa 5195 (3): 241-255, DOI: https://doi.org/10.11646/zootaxa.5195.3.3
