Cithaerias andromeda

Cithaerias andromeda (Fabricius)

(Figs 1a–c, 2, 6, 9a, 10)

Papilio andromeda Fabricius, 1775; Systema entomologiae, sistens insectorvm classes, ordines, genera, species, adiectis synonymis, locis, descriptionibvs, observationibvs. Flensburgi et Lipsiae, Korte

Type locality: “ India ”.

Type material: Likely lost (fide Lamas 1998). See item (1) in Historical overview.

Distribution. Venezuela eastward to French Guiana and southward to Brazil. See map (Fig. 10) for localities of examined specimens.

Diagnosis. This species is defined based on the following combination of characters (numbered in Fig. 2): (1) male HW submarginal band thin, usually separated from the marginal band and forming even, shallow arches; (2) male HW postmedial band that outlines the ocellus below M 1 usually complete across cells Rs through M 2, but it can be rather faded in some specimens; (3) male HW postmedial band forming even arches (thick or thin); (4) male HW scale cover blue-violet, diffuse, and usually extended into the discal cell, blue-violet scales outline veins M 2, M 3, CuA 1 and CuA 2; (5) female similar to male but with noticeably wider HW brown bands on the wing membrane; (6) female usually with reduced dorsal HW blue-violet scaling as compared to males, which is absent in some specimens. In males, FW brown bands vestigial to well-developed, extending from M 3 or CuA 1 to the anal margin. This band was present in all examined females. Male genitalia (Fig. 6): in lateral view, uncus straight; in dorsal view, the uncus and short tegument form an angular outline. Note variation between specimens of different localities in the width and inner sinuosity of the valva in ventral view. Saccus short and narrow anteriorly. Phallus straight, shorter than the length of the genitalic capsule in lateral view, and the vesica adorned with minute spines that extend slightly onto the shaft. Sub-anal plate triangular (posterior edge sometimes weakly sclerotized towards the midline), adjacent area membranous and bearing minute microtrichia, and the extension of microtrichia varies between localities. Examination of male genitalia revealed some variation between localities that match variation in hind wing color pattern (Fig. 6 shows genitalia of some of the specimens illustrated in Fig. 2). Female genitalia (Fig. 9a): sterigma tall and broad, V-shaped, post-sterigmal plate rounded, well-sclerotized and bearing minute microtrichia; ostium bursae narrow. Antrum short, and slightly more sclerotized than ductus bursae. Corpus bursae lacks signa. The female sterigma did not vary between localities within Guyana (Potaro River [CMP 13-32] and Mt. Ayanganna [CMP 19-09, not illustrated]).

Variation in wing pattern (Fig. 2). In males, the FW brown bands may be well-developed or reduced in both length and width (compare specimens in Fig. 2). HW postmedial, submarginal and marginal bands vary from particularly thin in some male specimens (Fig. 2d, i, j), to moderately thin (Fig. 2a, c, g), and wide in others (Fig. 2e). HW blue-violet scaling usually outlines wing veins (more prominent in males), but this is less obvious in worn specimens. In some localities, both males and females have pink scales. While HW white spots below veins M 2 and M 3 are always present in both sexes, a white spot or a complete ocellus may be present below CuA 1 (see Supernumerary ocelli section below). In females, the spread of blue-violet scaling varies from being almost as widespread as in the male sex (Fig. 2h), somewhat confined to cells M 3 and CuA 1 (Fig. 2f), or absent (Fig. 2b, k, l).

Subspecies. None at the present time. A study in progress is investigating divergence at the subspecific level (Mauro Costa Cicognani and collaborators, in prep.) through a more comprehensive geographical sampling than was possible here.

Remarks. The sparse blue-violet scale cover separates this species from other blue-colored taxa that have dense scaling. Given the variation in wing color and male genitalia (Fig. 2 and 6), identifying the type locality of this species is quite important. Lamas (1998) indicated that the type specimen of Fabricius’ Papilio andromeda likely originated from Suriname, and P. philis described by Stoll and illustrated in Cramer (1782, plate CCLXXXVII; Fig. 1a) is also from that country. Although the past cannot be recapitulated, it is possible that both Fabricius and Stoll published descriptions of the same taxon under different names during the seven years time span (1775–1782). Stoll’s illustration reproduced in Fig. 1a indeed matches specimens from Suriname (e.g., see species account and illustration in Gernaat et al. 2012), which likely justified the selection of a lectotype for P. phillis (type locality: Suriname) by Lamas (1998). Despite the lack of a type specimen associated with the nominal andromeda, this taxon can be identified with certainty because it is sufficiently distinct from other congeners.

Comparison of collection material to type specimen photographs (http://butterfliesofamerica.com/L/t/ Cithaerias _a.htm, last accessed December 2020) allowed me to assess the synonymy for C. andromeda proposed by Lamas (1998), which included five taxa (see Revised checklist for Cithaerias, above). Only one of the five type specimens is a male: the lectotype of P. philis (type locality: Suriname) designated by Lamas (1998). This specimen has a complete ocellus below CuA 1 that is not present in the illustration that accompanied the original description by Stoll (Fig. 1a), or in the illustration included in the Seitz catalogue (1910; Fig. 1c). In fact, the philis lectotype resembles the male in Fig. 2d (Guyana, “ Amazonas ”), including the pink spot within cell CuA 1 that is also present in some specimens from Venezuela (Fig. 2m). Although the holotype of C. philis f. hyacinthina (type locality: Óbidos, Brazil) is labeled as male, the broad FW and HW brown bands on the membrane suggest this specimen is a female of C. andromeda (part of the abdomen is missing). The holotypes of C. pellucida (type locality: French Guiana) and C. harpalyce (type locality: Upper Amazon?, see Historical overview, item (4)) are easily recognizable as females of C. andromeda, also strengthening the idea that early material was obtained from the former Dutch, British or French colonies (and not Brazil). Given the supernumerary ocelli, the female holotype of C. pellucida f. archeops is of particular interest and it is addressed below. Note that Mauro Costa Cicognani (pers. comm.) suspects that the collecting locality of the archeops type (San Fernando de Apure, Venezuela) might be inaccurate given various dubious records from Mayeul Grisol, named as the collector of the specimen on the collection label.

Supernumerary ocelli. The holotype of C. pellucida f. archeops is unusual as it has three faded ocelli on the FW (below R 5, M 1 and CuA 1) and three ocelli on the HW (a markedly large ocellus below M 1, a large ocellus below CuA 1, plus a faded one below CuA 2; http://butterfliesofamerica.com/L/ih/n_cithaerias0007_i.htm last accessed December 2020). This specimen has the largest number of supernumerary oceli of all Cithaerias specimens I have seen (including photographs), and its unusual appearance likely compelled Le Cerf to describe it as a form of C. pellucida. To my knowledge, C. andromeda is the only Cithaerias species in which supernumerary ocelli are not unusual and can be locally common. These were found to occur in specimens of multiple localities in French Guiana, Guyana, Brazil and Venezuela, as detailed below:

(1) French Guiana: The series of six specimens from Mana River (CMNH) includes one male where this ocellus is absent, but present in one female out of five (not illustrated). This ocellus is absent in a female from St. Jean Maroni (USNM, Fig. 2k) but clearly visible in the female from Maroni River (FMNH, Fig. 2l).

(2) Guyana: A small ocellus below CuA 1 is present in one of three females from Essequibo (FMNH) and a female from Potaro River (FMNH, Fig. 2b).

(3) Brazil: A pair from Pará, “Rio Tapajós” includes a female with a small ocellus below CuA 1, absent in the male (USNM, Fig. 2g –h). A male specimen from Amazonas, Manaus lacks this ocellus (FMNH), but it is present in a specimen from an adjacent locality (Reserva Florestal Adolpho Ducke, examined from a photograph provided by Leila Shirai). A photograph of one male and two females from Pará, Óbidos in the C. E. Mielke collection shows that the male lacks supernumerary oceli, but are present in the females (vestigial in one of them).

(4) Venezuela: In a pair from Bolívar, El Dorado (FMNH) the male lacks this ocellus, but it is present in the female (not illustrated). Mauro Costa Cicognani and Ángel Luiz Villoria Petit provided the following observations for Venezuela (pers. comm.): a large complete ocellus below CuA 1 was present in all specimens collected by Costa in a Venezuelan population from Sierra de Imataca (Fig. 2m, photographed with a flash). Costa believes that this population is homogeneous for the character, noting that it possibly represents local genetic differentiation. Villoria has not found C. andromeda with an extra HW ocellus outside the Imataca-Delta Amacuro region of Venezuela.

Finally, among all Cithaerias species and specimens I examined in a previous study (Penz et al. 2014), only a single specimen of C. aurora showed well-developed supernumerary ocelli where black rings are expressed around the white foci below veins Rs, M 2 and M 3, but not CuA 1 (Fig. 2n). In the early days of Cithaerias taxonomy, a specimen like this might have been described as a “form” of C. aurora.

Material examined: MALE: 1M, Brazil, Amazonas, Manaus, 10 Dec 1993 (FMNH); 1M, Brazil, Amazonas, Reserva Florestal Adolpho Ducke, 18 Jul 1985 (FMNH); 1♂, Brazil, Pará, Rio Tapajós, Aug 1983, 19-07 Dissected by CM Penz (USNM); 1M, Brazil, Pará, Santarém, 22 Aug 1985 (FMNH); 1M, Brazil, Paraná, Guarapuava, Jan 1983 (likely mislabeled; LACM); 1M, French Guiana, Cayenne (USNM); 1M, French Guiana, Mana River, Jun 1917 (CMNH); 1♂, French Guiana, Maroni River, 19-11 Dissected by CM Penz (FMNH); 20M, French Guiana, Maroni River (FMNH); 1M, British Guiana, Bartica (FMNH); 1♂, British Guiana, Holmia, Potaro Highlands, 8 Nov 1908, 19-10 Dissected by CM Penz (CMNH); 1♂, Guyana, Amazonas, Dec 1982, 19-13 Dissected by CM Penz (LACM); 1♂, Guyana, Berbice, New River Triangle, Camp Jaguar, + 500 m, 18 Nov 1980, 13-22 Dissected by CM Penz (FMNH); 1M, Guyana, Berbice, New River Triangle, Camp Jaguar, 18 Nov 1980 (FMNH); 1♂, Guy- ana, E bank Waruma River, 500m, 12 Mar–16 Apr 2001, 19-06 Dissected by CM Penz (USNM); 1♂, Guyana, Mt. Ayanganna, Kuiewa R., 2500–3300’, 2–25 Apr 1999, 19-05 Dissected by CM Penz (USNM); 1M, Guyana, N slope Mount Roraima, 1300 m, 12 Mar–16 Apr 2001 (USNM); 3M, Guyana, Potaro River, 29 Mar 2004, 4 Aug 2004 (FMNH); 1M, Venezuela, Bolivar, El Dorado, 5 Nov 1982 (FMNH). FEMALE: 1F, Brazil, Pará, Rio Tapajós, Aug 1983 (USNM); 1F, British Guiana, Kartabo Woods, 23 Jun 1925 (CMNH); 1♀, British Guiana, Potaro River, 29 Apr 1905, 13-32 Dissected by CM Penz (FMNH); 1F, Guyana, Bartica (FMNH); 2F, Guyana, 35 mi SW of Georgetown, Shanklands Resort, Essequibo River, 20–28 Sep 2006 (FMNH); 1F, Guyana, Essequibo, Lower Cuyuni River, 100 m, 29 Nov 1991 (FMNH); 1♀, Guyana, Mt.Ayanganna, 3300–4500’, 10–20 Apr 1999, 19-09 Dissected by CM Penz (USNM); 1F, Guyana, Potaro River, Nov–Dec 2002 (FMNH); 1F, French Guiana, near Caynne (sic), 4 May 1998 (FMNH); 1F, French Guiana, St. Jean Maroni (USNM); 5F, French Guiana, Mana River, Jun 1917 (CMNH); 18F, French Guiana, Maroni River (FMNH); 1F, Venezuela, Bolívar, El Dorado, 28 Nov 1984 (FMNH); 2F, Venezuela, Bolívar, La Escalera km 88 (San Isidro) 27 Jun 1984 (FMNH); 1F, no data (FMNH).