Calisto smintheus Bates, 1935

Aguila, Rayner Nunez, Plasencia, Edelquis Oliva, Maravi, Pavel F. Matos & Wahlberg, Niklas, 2012, Cuban Calisto (Lepidoptera, Nymphalidae, Satyrinae), a review based on morphological and DNA data, ZooKeys 165, pp. 57-105 : 66-70

Calisto smintheus Bates, 1935 stat. rev. Figs 4-62633414956, 576366-74

Calisto smintheus Bates 1935: 242

Calisto delos Bates 1935: 243, Michener 1943: 6, Schwartz and Hedges 1991: 136

Calisto smintheus smintheus Bates 1939: 3, Michener 1943: 6, Munroe 1950: 226, Torre 1952: 62, Torre 1954: 120, Torre 1968: 18, Núñez 2009: 56

Calisto smintheus delos Torre 1968: 19

Calisto biocellatus Torre 1968: 22

Calisto sibylla smintheus Brown and Heineman 1972: 51, Fontenla and Rodríguez 1990: 8, Smith et al. 1994: 57, Lamas 2004: 207

Calisto sibylla delos Brown and Heineman 1972: 51, Smith et al. 1994: 57, Lamas 2004: 207

Diagnosis.

Calisto smintheus requires comparison with some of its cogeners. Within Cuba, the more similar species is Calisto brochei ,but Calisto smintheus adults are larger on the average (19-25 mm of FWL versus 16-22 mm in Calisto brochei ), have a reddish suffusion around anal lobe at the UPHW, and are darker and more brightly colored at UN of wings. The androconial patch has a rounded outer margin in Calisto smintheus , but it is sinuous, forming three rounded lobes in Calisto brochei .Almost all other Cuban relatives except Calisto israeli , are paler and have fewer white dots at the post discal area on UNHW. Calisto herophile has also four white dots at UNHW, but is paler and smaller on the average, 14-21 mm versus 19-25 mm in Calisto smintheus . Outside Cuba, the Bahamian Calisto sibylla lacks red at the UNFW cell and the reddish suffusion at anal lobe; and in general, is a paler species. The Hispaniolan Calisto confusa Lathy, 1899, Calisto hysius (Godart [1824]) and Calisto obscura Michener, 1943, and Calisto pauli Johnson & Hedges, 1998 are superficially similar but all are distinctly smaller (13-18 mm) than Calisto smintheus .

Description.

FWL: 19-25 mm ♂ & ♀. Male UPFW dark brown except darker, almost black, androconial patch and postdiscal area adjacent to androconial patch and tornus, pale brown (Fig. 4). Androconial patch distinct except at base anterior limit, approximately triangular with outer margin rounded, anterior margin not entering into cell, about one half the length of FW (Fig. 33). Female UPFW dark grayish brown at basal two thirds, outer third pale grayish brown (Fig. 5). UPHW dark grayish brown at anterior two thirds, pale grayish brown at posterior third; anal lobe ferruginous, occupying apical half of posterior margin in some specimens. UN of wings brown heavily mixed with reddish and, toward base, pale yellow scales; apex of both wings and basal to pdl of HW with a dark wine hue (Figs 6, 26). Outer edge of pdl with bright yellow scaling. Post discal area at UNHW with four white dots at Rs–M 1, M1-M2, M2-M3, and M3-Cu1, the last one slightly displaced toward outer margin and smaller, sometimes absent in rubbed specimens. Male genitalia with tegumen about two thirds the length of uncus, rounded at posterior half (Fig. 41); uncus gradually tapering toward apex, arched at apical third; digitiform projection of valvae slender and long, straight at both margins; aedeagus sinuated with a left curve both at basal and apical half. Female genitalia with dorsal crown tall (Fig. 49); corpus bursae broad, about two thirds the length of ductus bursae.

Type material.

Holotype♂: Sierra del Cobre, Loma del Gato 3000 ft, 20°00'33"N, 76°02'16"W, 25-30/IX/1935, S. C. Bruner. MCZ, examined. Paratypes 8 ♂, 4 ♀: same data as for holotype except 2700-3300 ft, S. C. Bruner, genitalia ♂ & ♀ in glycerin. MCZ, CZACC, examined.

Calisto delos Bates 1935: holotype ♂, Ote (currently Santiago de Cuba), Pico Turquino, Loma Cordero (actually Cardero) 4000-6000 ft, 1 August 1935, J. Acuña; paratype ♂, Pico Turquino, Julio 22 de 1922, S. C. Bruner & C. H. Ballou, EEA Cuba No. 1652. MCZ, examined.

Calisto biocellatus Torre 1968: holotype ♂, Turquino, Pico Cuba 1872 m, 19°59'8.4"N, 76°50'32.3"W, VI/1963, F. de Zayas, P. Alayo & I. García; allotype ♀: same data as for holotype. CZACC, examined.

Additional material.

88 ♂, 33 ♀. Granma: Bartolomé Masó, La Platica 850 m, 20°00'54.1"N, 76°53'28.4"W, 26/XI/2007, R. Núñez, slide RNA175(androconial sclaes), DNA voucher PM07-05 (3 ♂); same data as for anterior except V/2008 (2 ♂). Santiago de Cuba: Aguada de Joaquín 1300 m, 20°00'50.4"N, 76°50'24.8"W, 20-27/I/2005, A. García, A. Barro & R. Núñez, genitalia ♂ in glycerin, slides RNA238(wings)/243(legs & labial palpus) (2 ♂, 1 ♀); same data as for anterior except 30/XI/2007, R. Núñez, genitalia ♀ in glycerin, slide RNA190(wings) (2 ♂, 1 ♀); Sierra Maetra, Pico Joaquín 5300 ft, 19°59'16"N, 76°53'31"W, 18/V/1948, J. Ferrás (3 ♂); ladera sur Pico Regino 1500 m, 20°00'38"N, 76°50'9"W, 29/XI/2007, R. Núñez, genitalia ♀ in glycerin, DNA voucher PM07-09 (M010) (1 ♂, 1 ♀); Sierra Maestra, 29/X/1941, J. Acuña (1 ♂); Turquino, June 1963, P. Alayo, slide RNA208(wings) (5 ♂); same data as for anterior except F. de Zayas, P. Alayo & I. García (1 ♀); Pico Turquino 1972 m, 19°59'23.7"N, 76°50'11.9"W, 18/X/1966, I. García, slide RNA275(legs & labial palpus) (10 ♂, 4 ♀); same data as for anterior except XII/1967, slides RNA225 (wings)/227(legs & labial palpus) (1 ♂, 1 ♀); same locality as for anterior, X/1985, M. G. Casanova, genitália ♀ in glycerin (1 ♂, 2 ♀); Ote (currently Santiago de Cuba), Turquino, Pico Cuba 1872 m, 19°59'8.4"N, 76°50'32.3"W, VI/1963, F. de Zayas, P. Alayo & I. García, genitalia ♂ & ♀ in glycerin, slides RNA186(androconial scales)/189/204/212(wings)/203/230/266 (legs & labial palpus) (10 ♂, 1 ♀); same locality as for anterior, 17/I/2002, A. Barro & R. Núñez (1 ♀); Ote (currently Santiago de Cuba), Sierra del Cobre, Loma El Gato 2600 ft, 20°00'33"N, 76°02'16"W, 24-30 September 1935, J. Acuña, S. C. Bruner & L. C. Scaramuzza (1 ♂, 1 ♀); same locality as for anterior, VIII/1942, Hno Crisogono (2 ♂); same locality as for anterior, 6/IX/1951, S. L. de la Torre, slide RNA228(wings) (8 ♂, 3 ♀); same locality as for anterior, 17-20 June 1952, F. de Zayas & P. Alayo (3 ♂); same locality as for anterior, 19 June 1952 (3 ♂); same locality as for anterior, 20 June 1952, slide RNA273(legs & labial palpus) (1 ♂); same locality as for anterior, 11/VIII/2008, E. Oliva, genitalia in glycerin, DNA vouchers PM07-13 (M030) & PM07-14 (M031) (2 ♂); same locality and date as for anterior, E. Fonseca (1 ♂); Ote (currently Santiago de Cuba), Caney, Gran Piedra 1100 m, 20°00'31"N, 75°37'3"W, Junio 1954, F. de Zayas & P. Alayo (1 ♀); same locality as for anterior, 23/IV/1955, P. Alayo, genitalia ♂ & ♀ in glycerin (2 ♂, 2 ♀); Ote (currently Santiago de Cuba), Caney, Gran Piedra, El Olimpo 900 m, 20°00'41"N, 75°39'42"W, 22 Mayo 1955, F. de Zayas & P. Alayo, slide RNA234(wings) (1 ♀); same data as for anterior except 26 Abril 1956, genitalia ♂ in glycerin, slides RNA192/221(wings) (4 ♂, 2 ♀); same data as for anterior except VIII/1960, genitalia ♀ in glycerin, slides RNA185(androconial scales)/188/219/251(wings)/216/276(legs & labial palpus) (8 ♂, 3 ♀); same locality as for anterior, VI/1962, P. Alayo, F. de Zayas & I. García (1 ♂); same locality as for anterior, 19/XII/1965 (4 ♂, 1 ♀); same locality as for anterior, 6/X/1966, I. García, genitalia ♀ in glycerin, slide RNA274(legs & labial palpus) (3 ♂, 3 ♀); same locality as for anterior, VIII/1986 (1 ♀); Gran Piedra, base Gran Piedra 1200 m, 16/III/2008, R. Núñez (4 ♂); Gran Piedra, pinar detrás Estación BIOECO 1100 m, 24/II/2011, R. Núñez (1 ♀); same data as anterior except ex ova, emerged 17/V/2011 (1 ♀). MFP, CZACC.

Distribution.

Species is restricted to the Sierra Maestra. It has been recorded from Pico Mogote ( Fontenla 2006) in the east to 140 km west at La Platica (Figs 56, 57). Besides anterior literature data, species has been recorded from La Bayamesa, Granma province ( Fontenla 2005).

Immature stages.

Egg & oviposition - Eggs are glued to substrate, spherical in shape and ivory white in color becoming beige with irregular orange brown spots a day after laid. Time to hatch 8 days (n=1).

First instar larva (Fig. 67) - Head capsule dark brown, almost black, with a bronze gloss and with two short horns on top. Body beige, greenish white on sides after fed on host leaves, with a dorsal line and four pairs of longitudinal pale orange brown lines: subdorsal, suprastigmatal, stigmatal, and infrastigmatal. Suprastigmatal line more greenish and the thinnest one, remainder lines more brownish and broader but subdorsal thinner than stigmatal and infrastigmatal lines. Dimensions (n=1): head capsule width 0.61 mm, head capsule height 0.64 mm, initial total length 2.9 mm, final total length 4.2 mm. Duration (n=1): 15 days.

Second instar with beige brown head capsule with slightly darker marks, body pattern similar to first but with a pair of dots, one at each subdorsal line, at metathorax that is present in remainder instars (Fig. 68). Instars from third and fourth with the same pattern of fifth, described below, but paler, with lines less contrasting, subdorsal and suprastigmatal lines straighter and the stigmatal and infragstigmatal lines distinct.

Fifth instar larva (Figs 69-71) - Head capsule beige regularly speckled with numerous dark brown dots; horns reduced; sides with two pairs of dark brown spots, each pair almost equidistant between them and to dorsal and ventral edges; mandibles black; X– mark of epicranium obsolete, represented only by a small rounded spot at apex of each arm, slightly darker than background. Body pale brown with brown striations; dorsum of each segment with darker “butterfly” like mark formed by small brown striations; lines slightly darker than background, except subdorsal which is pale yellow, lines becoming diffuse toward thorax; each abdominal segment with a transverse ashy gray band at beginning from dorsum to near suprastigmatal line and edged anteriorly by a brown dot at each end; dorsal line edged at beginning of each abdominal segment by two pale yellowish beige dots; a dark brown dot above subdorsal line on metathorax; subdorsal lines thinner than dorsal line, wavy, closer to dorsal line at middle of each segment, ending on caudal tails; suprastigmatal lines wavy following the wave pattern of subdorsal ones with dark brown dot above it near mid way to subdorsal, above it on each segment one pair of diffuse brown dots, one central, larger, and other near posterior margin; stigmatal and infrastigmatal lines diffuse mixed; area behind and below whitish, the latter crossed the infrastigmatal line. Dimensions (n=1): head capsule width 2.55 mm, head capsule height 2.58 mm, initial total length 14 mm, final total length 22 mm. Duration (n=1): 19 days.

Pupa (Figs 72-74) - Head and wing sheaths pale gray; antennae and leg sheaths with regular discontinuous pattern of dark brown dots; a pair of ventral black dots on eyes and another at sides of appendages near abdomen; wing sheaths edged at dorsum by an irregular dark brown large spot at middle; dorsum of thorax and abdomen pale gray with diffuse dark brown striations heavier at sides of dorsal ax forming a large spot on each side; abdomen with a dark brown line on sides, abdomen with a transverse ridge with a pair of more prominent crests on dorsum of segments 1 to 6; last abdominal segment long, stout, cremaster area enlarged, broad. Two days before emergence the dark brown extends covering almost entire thorax, extending gradually until occupying entire surface before emergence. Dimensions (n=1): total length 11 mm, maximum width 4.5 mm. Duration (n=1): 12 days.

Habitat and biology.

Throughout its range, the species inhabits evergreen and rainforests at altitudes between 800 m and 1500 m (Fig. 63). It is also found in cloud forest above 1500 m, and at the cloud scrub around Pico Turquino, 1972 m and Cuba highest peak. Individuals can be found in interior of forests but also at its edges. The species seems to prefer relatively well preserved areas but occasionally can be found at places with secondary vegetation. At La Platica village, Turquino massif, Sierra Maestra, the species was observed in shady places of gardens nearby forest, whereas, at Gran Piedra, it was found inside 25 year old pine plantations. Adults were observed feeding on flowers of Bourreria laevis , Palicourea alpina , Pavonia fruticosa , Mikania micrantha , and Stachyterpheta cayenensis in rainforest near La Platica.

Two females were observed when laid eggs singly at underside of leaves near midday. The host, Ichnanthus mayarensis , is the first one recorded for the Cuban species of the genus.This small grass is common at forest understory, sometimes abundant along paths, of rainforests in the Turquino Massif. Larvae eat the entire corion after hatching and feed at night remaining inactive during the day in lower parts of the plant. Larvae accepted both substitute host plants. First instar was 15 days long and all other were 9 days long each. Prepupal period was one day long and pupal stage extended for 12 days. Immature development takes 80 days and five larval instars.

Remarks.

Calisto smintheus and Calisto herophile are the only members of the genusinhabiting the Sierra Maestra. Their altitudinal ranges overlap between 800 and 1100 m, however, Calisto herophile is rare in places where Calisto smintheus is present and vice versa. Munroe (1950) mentioned the possibility of hybridization between them but there is no evidence available from present work to confirm it. The phylogenetic inferences and genetic distances agree on the establishment of Calisto smintheus as a single species with a minimum divergence of no lower than 5% from other Cuban Calisto taxa. The close phylogenetic relationship between Calisto smintheus and Calisto brochei is discussed below.