Cithaerias bandusia Staudinger, 2021

Penz, Carla M., 2021, Revised species definitions and nomenclature of the blue and purple / rose Cithaerias butterflies (Lepidoptera, Nymphalidae, Satyrinae), Zootaxa 4963 (2), pp. 293-316 : 302-303

Cithaerias bandusia Staudinger STAT. REV.

( Figs 3d–g , 7b , 9c , 10 )

Cithaerias esmeralda variety bandusia Staudinger, 1884 ; I. Theil. Exotische Tagfalter in systematischer Reihenfolge mit Berücksichtigung neuer Arten. In: Staudinger, O. & E. Schatz, 1884–1892, Exotische Schmetterlinge. Fürth, G. Löwensohn.

Type locality: Brazil, (Amazonas) .

Type material: The male lectotype designated by Lamas (1998) is deposited in the Museum Für Naturkunde , Berlin, Germany .

Distribution. Brazil, Amazonas, Mato Grosso, Pará. See map ( Fig. 10 ) for localities of examined specimens.

Diagnosis. This species is defined based on the following combination of characters (numbered in Fig. 3d–g ): (1) male HW submarginal and marginal bands amalgamated and thin; (2) male HW postmedial band proximal to the ocellus below M 1 thin, fragmented, often limited to cell Rs; (3) male HW postmedial band quite broad around tornus, may be absent from cell M 3; (4) male HW scale cover blue, glossy, dense and limited to the tornus, covering the postmedial band and enclosing medium-small pink spots; (5) female similar to male but with wider HW brown bands, a continuous postmedial band including the portion outlining the ocellus below M 1, (6) less dense midnightblue scaling at tornus enclosing larger pale pink or white spots. Male genitalia ( Fig. 7b ): in lateral view, uncus bent near base; in dorsal view, the uncus and short tegument form a shallowly sinuous outline. In ventral view, valva relatively thin, sinuous. Saccus broad, triangular-shaped, longer than in esmeralda . Phallus long and sharply bent at approximately 2/3 the length from the base; phallus bend narrower than adjacent points of the shaft. Sub-anal plate somewhat rectangular posteriorly and narrowing anteriorly, adjacent area mildly sclerotized with larger microtrichia than esmeralda . Female genitalia ( Fig. 9c ): sterigma short, somewhat rectangular; ostium bursae wide. Post-sterigmal area weakly sclerotized, rounded, bearing minute microtrichia. Antrum long, slightly sclerotized only near the ostium bursae, ductus bursae bulging anteriorly (unique to this species); the bulge in the ductus bursae and length of the antrum plus ductus bursae seem to correspond to the morphology of the male phallus (bend and length). Corpus bursae lacks signa.

Variation in wing pattern ( Fig. 3d–g ). Males and females show some variation in the size of the HW pale pink spots enclosed within the blue scaling at tornus. The brown bands in the HW membrane can be markedly reduced in some male specimens.

Subspecies. None.

Remarks. Both male and female C. bandusia show remarkable differences in color pattern and genitalia morphology from C. andromeda where they had been placed by Lamas (1998) These differences include: dense hind wing scaling, elongate and bent phallus, bent uncus, and a short and delicate sterigma (compare Fig. 3d–g to Fig. 2 , Fig. 7b to Fig. 6 , Fig. 9c to 9a). Moreover, C. bandusia has been collected at the same locality with C. esmeralda ; i.e., Brazil, Pará, Santarém. This species differs from C. esmeralda by having blue hind wing color (compare Fig. 3d–g to 3a–c) and a sharply bent phallus, which is evenly arched in C. esmeralda , (compare Fig. 7b to 7a). Although pairs have not been examined in copula, the bend in the phallus ( Fig. 7b ) correlates with a bulge in the female ductus bursae ( Fig. 9c ), suggesting the concerted evolution of male and female genitalia morphology. These characters are absent in C. esmeralda ( Fig. 7a , 9b ). Thus, the species status for C. bandusia proposed here is justified by wing color and genitalia characters plus putative sympatry with C. esmeralda . Note that Santarém is the easternmost locality for C. bandusia recorded in the collections examined here, and that this species has also been apparently collected in Óbidos (Pará, Brazil), which is located on the northern bank of the Amazon River.

Behavior. The following field observations were provided by Phil DeVries (pers. comm., written by him and included here with permission). “During June 2009 (dry season) I found that C. bandusia was moderately abundant in the flood plane forest at Cristalino Lodge ( Brazil, Mato Grosso, Alta Floresta; 9.8672 o S, 56.0870 o W). Based on my experience with other Cithaerias species , individuals of C. bandusia behaved as I had anticipated; throughout the day they glided just above the forest floor through the understory, along trails and in light gaps, perching on low vegetation ( Fig. 4 ) and sometimes interacting as pairs. What was interesting and noticeably different to me was their behavior in the late afternoon about one hour before sunset. When the sunlight was streaming nearly horizontally through the forest I noticed that individual butterflies were, by and large, facing west, slowly and repeatedly elevating and dropping their open hind wings into the light. At certain angles of observation this behavior produced small iridescent blue flashes. This was most evident when lying down on the forest floor on my stomach approximating the level of the butterflies themselves, and looking at a slightly WNW direction down a wide central trail. At times there were multiple individuals producing these flashes. This suggested to me that perhaps C. bandusia individuals were signaling to each other in a call and answer fashion like ships at sea using signal lamps to communicate. I observed this phenomenon over the course of about a week, always late in the day when the sun was on the horizon. Although I never saw courtship or mating, these simple field observations suggest that some potentially important behaviors of Cithaerias are confined to very short periods of the day.”

Material examined: MALE: 2M, Brazil, Amazonas, Maués , Nov 1988 ( FMNH) ; 1M, Brazil, Mato Grosso, Alta Floresta , 23 May 2009 ( PJD) ; 1M, Brazil, Luré , Mar 1987 ( FMNH) ; 1M, Brazil, Pará, Itaituba, R. Tapajós, Jul ( FMNH); 3M, Brazil, Pará, Óbidos , Feb–Mar 1989, Jan 1988 ( FMNH) ; 1M, Brazil, Pará, Óbidos , Mar 1984 ( LACM) ; 1♂, Brazil, Pará, Rio Tapajós , 8 Aug 1983, 19-08 Dissected by CM Penz ( USNM); 1M, Brazil, Pará, Rio Tocantins, Alcobaça (sic) ( ETHZ) ; 2M, Brazil, Pará, Santarém , Feb 1984, 10 Oct 1984 ( FMNH) ; 2M, Brazil, Pará, Santarém , 14 Feb 1984, 11 Feb 1984 ( FMNH) ; 1♂, Brazil, Pará, Santarém , Nov 1988, 14-10 Dissected by CM Penz ( LACM); 1M, Brazil, Pará , Jul 1984 ( FMNH) ; 1M, French Guiana, Commune de Roura , Jan 1998 (likely mislabeled, FMNH) ; 1M, French Guiana, Commune de Roura , Mar 1991 (likely mislabeled, FMNH) ; 1M, Brazil, Joinville , 14 Dec 1986 (clearly mislabeled, FMNH) ; 1M, no locality ( CMNH). FEMALE: 1F, Brazil, Amazonas, Maués , Dec 1922 ( FMNH) ; 5F, Brazil, Pará, Óbidos , Feb–Mar 1989, 9 Jun 1985 ( FMNH) ; 1F, Brazil, Pará, Rio Tapajós , 8 Aug 1983 ( USNM) ; 1F, Brazil, Pará, Rio Tocantins, Alcobaça (sic) ( ETHZ) ; 1F, Brazil, Pará, Santarém , 28 May 1987 ( FMNH) ; 1♀, Brazil, Pará, Santarém , 14 Oct 1984, 14-11 Dissected by CM Penz ( LACM); 1F, Brazil, Pará , 1885 ( CMNH) ; 1F, French Guiana, Commune de Roura , Mar 1996 (likely mislabeled, FMNH) .

Figures

FIGURE 2. a–m, C. andromeda; composite pictures show dorsal view on the left, ventral on the right. n, C. aurora (C. Felder & R. Felder). Collecting localities and genitalia dissection codes appear below each specimen. Mauro Costa Cicognani provided the photograph of one of the C. andromeda specimens (m), and granted permission for its use here.

FIGURE 3. a–c, C. esmeralda; d–g, C. bandusia; and h–i C. azurina; dorsal view on the left, ventral on the right. Collecting localities and genitalia dissection codes appear below each specimen. Cithaerias azurina specimens were photographed before dissection, collection labels were reduced to 70% of their original size.

FIGURE 4. A male C. bandusia from Cristalino Lodge (Brazil, Mato Grosso, Alta Floresta; June 2009). Photo by Phil DeVries, used here with permission. A specimen from this locality was included in the phylogeny by Matos-Maraví et al. (2019).

FIGURE 6. Male genitalia in lateral and ventral views, tegumen and uncus in dorsal view, phallus in lateral view, and subscaphium in ventral view. Valva setae shown on one side only. Given that the two parts of the subscaphium (subanal plate and adjacent area containing microtrichia) rest on the diaphragma, the variation in the shape of the subanal plate might include slight preparation artifacts. Scale bar 1 mm for all images. a, C. andromeda (Guyana, Berbice) dissection CMP 13-22; b, C. andromeda (Guyana, Mt. Ayanganna) dissection CMP 19-05; c, C. andromeda (French Guiana, Maroni River) dissection CMP 19-11.

FIGURE 7. Male genitalia in lateral and ventral views, tegumen and uncus in dorsal view, phallus in lateral view, and subscaphium in ventral view. Valva setae shown on one side only. Given that the two parts of the subscaphium (subanal plate and adjacent area containing microtrichia) rest on the diaphragma, the variation in the shape of the subanal plate might include slight preparation artifacts. Scale bar 1 mm for all images. a, C. esmeralda (Brazil, Pará) dissection CMP 19-03; b, C. bandusia (Brazil, Pará, Rio Tapajós) dissection CMP 19-08; c, C. azurina ([Brazil, Amazonas] Rio Papori (sic)) dissection CMP 19-14.

FIGURE 9. Female genitalia in ventral view, including schematic representation of papilla anales and abdomen outline (scale patches, general scaling, and setae excluded). Scale bar 1 mm for all images. a, C. andromeda (British Guiana, Potaro River) dissection CMP 13–32; b, C. esmeralda (Brazil, Pará) dissection CMP 19-04; c, C. bandusia (Brazil, Pará, Santarém) dissection CMP 14-11; d, C. azurina ([Brazil, Amazonas] Rio Papori (sic)) dissection CMP 19-15; e, C. pyropina pyropina (Peru, Tingo María) dissection CMP 13-31; f, C. pyropina julia (Ecuador, Macas) dissection CMP 19-16; g, C. songoana (Bolivia, Rio Songo (sic)) dissection CMP 20-17. The arrow in e shows a separate drawing of the sclerotized post-sterigmal plate (folded behind the sterigma in the prepared specimen, and also in f and g). This structure is present in all species examined, and all have microtrichia, which vary in size between species.

FIGURE 10. Map showing the collecting localities of examined specimens (excluding dubious localities). Circles marked with a white dot represent specimens examined from photographs.