Amiga arnaca arnaca (Fabricius, 1776) Fabricius, 1776

Nakahara, Shinichi, Lamas, Gerardo, Tyler, Stephanie, Marin, Mario Alejandro, Huertas, Blanca, Willmott, Keith R., Mielke, Olaf H. H. & Espeland, Marianne, 2019, A revision of the new genus Amiga Nakahara, Willmott & Espeland, gen. n., described for Papilioarnaca Fabricius, 1776 (Lepidoptera, Nymphalidae, Satyrinae), ZooKeys 821, pp. 85-152 : 94-99

Amiga arnaca arnaca (Fabricius, 1776) comb. n. Figs 2 a–h, 3, 4 a–d, 4 f–k, 5, 6

Papilio arnaca Fabricius (1776: 260-261). Type locality: Suriname. Neotype ♂ (here designated): // Suriname Brokopondo Brownsberg, rainforest km 6-12, 30.1.1982, Olle Pellmyr // USNMENT 00913953 // (USNM) [examined]

= Papilio ebusa Cramer (1780: 9; pl. CCXCII: figs F, G). Type locality: Suriname. Neotype ♂ (here designated): Suriname Brokopondo Brownsberg, rainforest km 6-12, 30.1.1982, Olle Pellmyr // USNMENT 00913953 // (USNM) [examined]

= Euptychia arnaea [sic] form priamis D’Almeida (1922: 99). Type locality: Três Rios, Jacarepaguá [Rio de Janeiro (city), Rio de Janeiro (state), Brazil]. Holotype ♂: // Euptychia ar-naea priamis d’Alm. 1922. ♂ // Jacarépaguá, Tres-Rios. Rio, 19.. ♂ // HOLOTIPO// Coll. D’Almeida // No 5630 // DZ 34.684// (DZUP) [examined]

Papilio arnaea [sic]: Fabricius 1781: 85; Fabricius 1787: 37; Kirby 1871: 53; Butler and Druce 1874: 337; Butler 1876: 489; Kaye 1904: 181; Hall 1939: 34.

Papilio aranea [sic]: Fabricius 1793: 97.

Satyrus aranea [sic]: Godart [1824]: 492; Butler 1867: 489.

Euptychia ebusa : Butler 1867: 489; Möschler 1877: 323; Kirby 1879: 135; Godman and Salvin 1880-1881: 88-89; Dyar 1914: 143.

Euptychia arnaea [sic]: Kirby 1871: 53; Butler and Druce 1874: 337; Butler 1877: 122; Möschler 1877: 323; Godman and Salvin 1880-1881: 88-89; Sharpe 1890: 569; Kaye 1904: 181; Weymer 1911: 219, pl. 49d; Hall 1939: 34; Whittaker 1983 (misspelled as “arneae” in figs 1, 2); D’Abrera 1988: 770-771; Emmel and Austin 1990: 10; Mielke and Casagrande 1991: 181; Cock 2014: 11.

Euptychia arnaea [sic] form priamis : D’Almeida 1937: 254; Brown 1975: 41.

Euptychia arnea [sic]: Gaede 1931: 439; Barcant 1970: 143, pl. 13, fig. 18.

Euptychia arnea [sic] var. priamis : Gaede 1931: 439.

Euptychia arnaca arnaca : Bryk 1953: 63.

Chloreuptychia arnea [sic]: Forster 1964: 120, fig. 131.

Chloreuptychia arnaea [sic]: DeVries 1987:271, pl. 48 figs 18, 19; 261, figs B, C; Ramos 1996: 40; Brown and Freitas 2000: 105.

Cissia arnaea [sic]: Singer and Ehrlich 1993 251, fig. 1.

Chloreuptychia arnaca : Lamas 1994: 165; Lamas and Grados 1996: 58; Lamas et al. 1996: 65; Lamas et al. 1999: 10; Lamas 2004: 218; Beccaloni et al. 2008: 328; Brévignon 2008: 70; Marín and Uribe 2009: 24; Peña et al. 2010: 246; Francini et al. 2011: 65; Paluch et al. 2011: 235; Brévignon and Benmesbah 2012: 52; Cock 2014: 11; Freitas et al. 2016: 320; Paluch et al. 2016: 4.

Identification and taxonomy.

Papilio arnaca Fabricius, 1776 was described based on an unspecified number of specimens from Suriname, in Johann Dominicus Schulze's collection ( Fabricius 1776). Fabricius' description was not accompanied by any illustration of this species, and he did not specify either the sex nor the number of specimens he examined. However, his Latin description is somewhat precise and the identity of the species may be guessed from the description, given the mention of the following wing pattern characters: "forewing, towards the apex there are three ocelli: the distalmost ("exteriori") bi-pupilled"; "hindwings bluish; under surface with five ocelli"; "Hindwings bluish above, iridescent; below bluish with two oblique dark stripes. Submargin with five ocelli, the first and fourth the largest and black, the remainder dark". The mention of multiple ocelli on the (ventral) forewing excludes the possibility of this specimen being other " Chloreuptychia " species, which also possess bluish iridescent coloration, but have only a single ocellus on the ventral forewing. Fabricius (1793) considered P. lea Cramer, 1777 and P. arnaca as probably being conspecific, and these two names were associated by some subsequent authors (e.g., God man and Salvin 1880-1881), although the mention of five ventral hindwing ocelli for P. arnaca is inconsistent with the six ocelli on the ventral hindwing of P. lea . On the other hand, Neonympha iris C. Felder & R. Felder, 1867 and Euptychia tricolor Hewitson, 1850, two species now placed in Magneuptychia Forster, 1964, also match the aforementioned wing pattern characters provided in Fabricius’ original description. These two species do occur in Suriname, and based on the description provided for P. arnaca , it is difficult to exclude the possibility of Fabricius having examined one of these two species. Regardless of this fact, the name arnaca has been applied in numerous publications and collections to the species as it is identified here (e.g., Whittaker 1983). Considering this situation, stabilizing the nomenclature as currently perceived by many others is crucial regarding the specific epithet arnaca and a neotype is therefore designated for this name below.

A worn specimen (whose sex cannot be confidently determined) in William Hunter’s entomological collection is at the UMG, and it was photographed by GL as a potential type specimen of P. arnaca (see Warren et al. 2018). GL assumed that William Hunter may have received a "duplicate" from Schulze through Fabricius, who did in deed supply Hunter with duplicates, resulting in many Fabrician type specimens being found in Hunter’s insect collection ( Hancock 2015; Tuxen 1967). However, we have found no evidence to support that this specimen was originally in Johann Dominicus Schulze's collection, and the Surinamese provenance of the specimen is questionable given its rather narrow ventral bands, which are typical of A. arnaca indianacristoi ssp. n. rather than of specimens from Suriname. The specimen in Glasgow is missing its head and abdomen, in addition to having worn and faded wings, thus somewhat obscuring its true identity. Given this situation, combined with the fact that no authentic Schulze specimens appear to be in existence (e.g., Benmesbah et al. 2018), in addition to the explanation above, we therefore designate a male specimen from Suriname (type locality) as a neotype for P. arnaca following Article 75.3 of the ICZN (1999) (neotype designation): //Suriname Brokopondo Brownsberg, rainforest km 6-12, 30.1.1982, Olle Pellmyr // USNMENT 00913953// (USNM). (Fig. 2a).

After introducing this species to science, the specific epithet was misspelled as “Arnaea” by Fabricius himself in 1781 and 1787, and as “Aranea” in 1793. Subsequently, the specific epithet arnaca has been erroneously spelled in various ways in a disturbingly high number of publications (e.g., Butler and Druce 1874; Butler 1877; Godman and Salvin 1881; Sharpe 1890; Kaye 1904; Weymer 1911; D’Almeida 1922; Gaede 1931; Forster 1964; Brown 1975; Whittaker 1983; DeVries 1987; see also above), including some influential works on the classification of this group. This confusion surrounding its species-group name adds a special urgency for a neotype designation for this common butterfly.

Papilio ebusa Cramer, 1780 was described in Pieter Cramer’s De uitlandsche Kapellen voorkomende in de drie Waereld-Deelen Asia, Africa en America. The original description describes the bluish-lilac reflection on both wing surfaces, although no further description of any wing element was provided in Cramer’s Dutch and French description. Instead, Cramer compared P. ebusa to P. junia Cramer, 1780, an immediately preceding species described and named in Cramer (1780), but regarded as a junior subjective synonym of P. lea Cramer, 1777 by Lamas (2004). Evidently, P. ebusa and P. junia are not conspecific judging from the illustrations in Cramer (1780: 9; pl. CCXCII: figs D–G), and the illustrations of P. ebusa combined with Cramer’s description enable this taxon to be confidently identified. Papilio chloris Cramer, 1780 (now known as Chloreuptychia chlorimene ( Hübner, [1819])), is perhaps the only taxon known from Suriname which might have resulted in a similar illustration; however, the illustration of P. chloris provided by Cramer (1780: CCXCIII: figs A, B) excludes this possibility. Based on the Dutch and French description provided for P. ebusa , Cramer based his illustration on what he thought was a female specimen, although the illustration of the dorsal surface (Fig. F) showing the bluish-lilac reflection only on the hindwing indicates that this illustrated specimen is likely to be a male (but see also above for further information). In addition, whether the original description was based on a single specimen or several specimens cannot be unambiguously determined. During our attempt to locate syntype(s) of P. ebusa , two specimens with rounded labels indicating "[Johan] Calkoen" with the locality “Brasilia” were found in RMNH. Along with the collection of Joan Raye Heer van Breukelerwaard, Johan Calkoen’s collection includes Cramer types, although given the locality “Brasilia”, these two specimens are most likely not syntypes of P. ebusa . Considering that we were unable to find any additional possible syntype(s) of P. ebusa , we here designate a neotype for this name. Although treated as a valid species in the past (e.g., Butler 1867; Godman and Salvin 1880-1881), in order to maintain its status as a junior synonym of P. arnaca , first recognized by Kirby (1871) and followed by most subsequent authors (e.g., Weymer 1911; Gaede 1931; Lamas 2004), we designate the specimen designated as the neotype of P. arnaca as the neotype of P. ebusa as well and retain its synonymy as a junior objective synonym (neotype designation).

D’Almeida (1922) described Euptychia arnaea [sic] form priamis based on a single male from Três Rios, Jacarepaguá, Rio de Janeiro, Rio de Janeiro, Brazil, currently housed at the DZUP. Following Article 73.1.2. of the ICZN (1999), we consider this male specimen to be the holotype fixed by monotypy based on the statement of "one male collected at the type locality" provided in the original description ( D’Almeida 1922). Lamas (2004) regarded this taxon as a junior subjective synonym of Papilio arnaca without providing any justification. D’Almeida’s (1922) original description provides some wing pattern characters which he considered to separate f. priamis , namely "Underside, feeble pearly reflections extending from the base to the line of ocelli"; "Underside, the two rays in the middle are narrow". These two wing pattern characters are seen in the holotype male, and indeed, the overall phenotype of specimens from the Brazilian states of Minas Gerais, Espírito Santo, and Rio de Janeiro does look somewhat different compared to the neighbouring nominotypical subspecies. Although the feeble pearly reflection extending from the base of the ventral forewing is not seen in the nominotypical subspecies, a few specimens from the aforementioned states in southeastern Brazil appear to lack this reflection (e.g., FLMNH-MGCL-1036218). The narrow ventral bands of many specimens from southeastern Brazil resemble those of A. arnaca indianacristoi ssp. n., although the ventral bands are slightly variable in width and a few specimens (e.g., FLMNH-MGCL-262982, 263014) possess bands that are similar in width to the nominotypical subspecies. Thus, the majority of the specimens from Minas Gerais, Espírito Santo, and Rio de Janeiro are distinguishable from the nominotypical subspecies based on the aforementioned characters except for specimens from Bahia consistently possessing wider ventral bands and/or lacking the feeble pearly reflections on the ventral surface. Nevertheless, we decided not to treat A. arnaca from Minas Gerais, Espírito Santo, and Rio de Janeiro as a distinct subspecies because, based on molecular data, this taxonomy would result in the nominotypical subspecies being paraphyletic. Whether subspecies should simply represent geographical variation or should also represent an evolutionary unit (i.e. a monophyletic group) is not a focus of this study and this question merits further in-depth discussion and more data. To be consistent in terms of the subspecies concept used in this study, we consider that subspecies should ideally represent clades, unless there is a strong counter-argument, and thus retain the synonymy introduced in Lamas (2004).

Distribution

(Fig. 6). The nominotypical subspecies occurs from eastern Colombia south to Bolivia, and in Brazil, southern Venezuela and the Guianas, where it is typically common and widespread in lowland to submontane forest.

Examined specimens

(718 ♂, 207 ♀). See Appendix for the data of these specimens.