Calisto herophile Huebner , 1823

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 : 82-88

Calisto herophile Huebner, 1823 Figs 22 –243139475556–596690– 99

Calisto herophile Hübner 1823: 16, Gundlach 1881: 111, Lathy 1899: 223, Dethier 1940: 14

Satyrus herophile Poey, 1847: 179

Calisto herophile herophile Bates 1935: 242, Michener 1943: 6, Michener 1949: 1, Munroe 1950: 225, Torre 1952: 62, Torre 1954: 120, Torre 1968: 12, Brown and Heineman 1972: 51, Alayo and Hernández 1987: 39, Schwartz and Hedges 1991: 136, Smith et al. 1994: 56, Lamas 2004: 2007, Núñez 2009: 56

Diagnosis.

Calisto herophile can be separated by its similar congeners in several ways. From Calisto smintheus and Calisto brochei ,it differs, among other features, by its paler background color at both sides of wings, the inconspicuousness of its androconial patch and its less sclerotized male genitalia with a shorter uncus and less sinuous aedeagus. From Calisto muripetens , Calisto occulta and Calisto bradleyi ,it differs by having four white dots and paler coloration. Differences with Calisto bruneri are detailed in the Diagnosis section of that species. It is also similar the Bahamian Calisto sibylla but smaller, 14-21 mm of FWL versus 23 mm in Calisto sibylla which also lacks the red in cell at the UNFW present in Calisto herophile . The Hispaniolan Calisto confusa , Calisto hysius and Calisto obscura although similar in size are darker,and have straighter white edged lines at the UNHW. Other Hispaniolan species, Calisto pauli , is similar in size and pattern but has different genitalia including a larger and flattened uncus in males and a terminal production in the dorsal crown of the female genitalia.

Description.

FWL: 14-19 mm ♂, 17-21 mm ♀. Male UP of wings dark brown at basal area more or less defined by UN pdl, area outer to pdl distinctly paler (Fig. 22). Androconial patch indistinct in fresh specimens, approximately triangular with apex slightly angled, anterior margin not surpassing posterior margin of cell, about two fifths the length of FW (Fig. 39). Female UP of wings as in male but distinctly paler (Fig. 23). UNHW background pale brown heavily mixed with pale yellow scales (Figs 24, 31). Post discal area on UNHW with four two white dots at Rs–M 1, M1-M2, M2-M3, M3-Cu1 interspaces, the last one, and occasionally the first one too, smaller and sometimes absent in rubbed specimens. Male genitalia with tegumen about two thirds the length of uncus, nearly straight, posterior end rounded (Fig. 47); uncus broad at basal half, tapering gradually from the middle toward apex, arched at apical third; digitiform projection of valvae with ventral margin straight; aedeagus only slightly sinuated in dorsal view, with two small left curves at apical half. Female genitalia with dorsal crown tall (Fig. 55); corpus bursae somewhat broad, about 0.6 the length of ductus bursae.

Type material.

Holotype♂: Cuba, Havannah. Location unknown, not examined.

Additional material.

148 ♂, 76 ♀. Pinar del Río: Pinares de Viñales 200 m, 22°35'N, 82°42'41"W, V/1963, P. Alayo & I. García, genitalia in glycerin, slide RNA 223(legs & labial palpus) (1 ♂); Rangel 400 m, 22°45'N, 83°11'W, 2/XI/1966, I. García & S. L. de la Torre (4 ♂); same data as for anterior except I. García, slide RNA196(wings) (1 ♀); same locality and collector as for anterior 21/VII/1967 (9 ♂, 6 ♀); same locality as for anterior, R. Núñez & E. Oliva, 19-20/IV/2009, ex ova, emerged 19/VI/2009 (2 ♂); same data as for anterior except emerged 20/VI/2009 (1 ♀); 22/VI/2009 (1 ♂); 23/VI/2009 (1 ♀); 26/VI/2009 (1 ♂); Viñales 150 m, 22°36'59"N, 82°42'28"W, 21/VII/1967 (6 ♂); same locality as for anterior, X/1985, J. L. Fontenla (2 ♂); Valle de Viñales, 9/I/1974, A. Castiñeiras (1 ♂); carretera a Viñales km 22 200 m, 22°34'29"N, 82°42'11"W, 14/I/1974, A. Castiñeiras (1 ♀). Mayabeque: Jaruco, Cueva Don Martin, 23°00'N, 82°01'W, 4/V/1966 (5 ♂); La Habana (currently Artemisa), Guajaibón próximo a Mariel, 23°01'N, 82°40'52"W, 25/V/1967 (1 ♂, 1 ♀); same locality as for anterior, X/2007, R. Núñez, DNA voucher PM07-07 (M008) (1 ♀); Pinar del Río (currently Artemisa), Sierra del Rosario, III/1968, R. González (1 ♂); Pinar del Río (currently Artemisa), Sierra del Rosario, alrededores Estación Biológica 180 m, 22°51'N, 82°55'53"W, 1-10/X/2007, R. Núñez (1 ♂); Sierra del Rosario, El Mulo 200 m, 22°51'29"N, 82°56'54"W, 10/X/2007, R. Núñez (1 ♂). Isla de La Juventud: Isla de Pinos (currently Isla de La Juventud), Cerro San Pedro 150 m, 21°42'47"N, 82°51'50"W, 20/X/1966, I. García (2 ♂); Habana (currently Isla de La Juventud), Isla de Pinos (currently Isla de La Juventud), 30/X/1966, I. García (7 ♂, 7 ♀). Habana: Cerro, 23°06'27"N, 82°23'20"W, 9/I/1934 (1 ♂); Arroyo Naranjo, 23°01'N, 82°22'W, 5 August 1935, L. C. Scaramuzza (1 ♀); Santiago de Las Vegas, 22°58'N, 82°23'W, 15/VIII/1935, S. C. Bruner, genitalia in glycerin (1 ♀); same locality as for anterior, 5 Marzo 1946, J. Ferrás (1 ♂); same data as for anterior except 19 March 1948 (2 ♂); Cotorro, 23°02'N, 82°16'W, 1/XII/1947, J. T. Sierra (1 ♂). Mayabeque: Matanzas (currently Mayabeque), 5 km W Ceiba Mocha 150 m, 22°58'50"N, 81°46'24"W, 8/IX/1940, S. L. de la Torre (1 ♂); La Habana (currently Mayabeque), Madruga, La Jiquima 125 m, 22°53'58"N, 81°50'34"W, 5/X/1948, S. L. de la Torre & J. Ortiz (1 ♀). Matanzas: Los Prácticos, 23°02'37"N, 81°34'32"W, 23/VII/1940, S. L. de la Torre (1 ♂); Playa, 23°02'37"N, 81°34'32"W, 11/V/1942, S. L. de la Torre (1 ♀); same data as for anterior except 16/VI/1942, slide RNA242(wings) (1 ♂); same data as for anterior except 29/VIII/1947 (1 ♀); same data as for anterior except 6/X/1947 (1 ♂); same data as for anterior except 26/VIII/1948 (1 ♀); same data as for anterior except 27/VIII/1948 (1 ♂); same data as for anterior except 6/XI/1948 (2 ♂); km 6 Vía Blanca, Playa Mamey, 23°03'06"N, 81°29'41"W, 6/VII/1953, S. L. de la Torre (1 ♀); Varadero, Varahicacos, 23°11'40"N, 81°09'16"W, 17/VI/2008, R. Núñez, slide RNA218(wings), DNA voucher PM15-04 (2 ♂). Cienfuegos: Las Villas (currently Cienfuegos), Escambray, Mina Carlota 450 m, 22°03'55"N, 80°09'38"W, 15/VI/1967, genitalia ♂ & ♀ in glycerin, slides RNA207(legs & labial palpus)/206(wings) (3 ♂, 2 ♀); Las Villas (currently Cienfuegos), Escambray, Buenos Aires 700 m, 21°59'13"N, 80°11'20"W, 16/VI/1967, genitalia ♂ & ♀ in glycerin, slides RNA182(androconial scales)/203(legs & labial palpus)/226/232(wings) (9 ♂, 4 ♀); Escambray, Charco Hediondo a 10 km de Aguacate, VIII/1978, L. R. Hernández (1 ♂). Villa Clara: Mordazo, 22°38'29"N, 80°26'58"W, V/1934 (1 ♀). Sancti Spiritus: Trinidad, La Vigía 200 m, 21°48'48"N, 79°58'34"W, 15/VI/1967 (1 ♂). Camagüey: Camagüey, 21°22'51"N, 77°55'01"W, 23/IX/1967, S. L. de la Torre (6 ♂). Holguín: Ote (currently Holguín), Pinares de Mayarí 800 m, 20°28'8"N, 75°48'52"W, 16/X/1966, I. García (10 ♂, 5 ♀); same locality as for anterior, VI/1967, P. Alayo (1 ♂, 2 ♀); Moa, El Johnson 300 m, 20°35'36.4"N, 74°59'9.9"W, 5/I/1968, S. L. de la Torre, slide RNA 167(wings) (1 ♂); same data as for anterior except 6/I/1968 (1 ♂); Moa, Quemado del Negro, 22°36'40"N, 74°49'22"W, 6/I/1968, S. L. de la Torre (1 ♂, 1 ♀); same data as for anterior except 7/I/1968, slide RNA281(legs & labial palpus) (3 ♂, 3 ♀); Mayarí, camino de La Zoilita 250 m, 20°38'N, 75°29'W, IX/1986, R. Rodríguez, genitalia in glycerin (2 ♂); Mayarí, El Purio, 20°39'45"N, 75°30'55"W, IX/1986, R. Rodríguez, genitalia ♀ in glicerin, slide RNA220(wings) (2 ♂, 2 ♀); Jaguaní, Arroyo Bueno o La Melba 200 m, 20°26'24"N, 74°48'46"W, VIII/2001, R. Núñez (1 ♂, 1 ♀); antiguo campamento minero Meseta de El Toldo 815 m, 20°27'35"N, 74°53'53"W, V/2008, E. Pérez (3 ♂); Moa, km 1 camino de La Melba, 20°36'12"N, 74°50'20"W, 19/I/2009, R. Núñez, genitalia ♀ in glycerin, slide RNA259(legs & labial palpus), DNA voucher PM15-06 (M052) (1 ♂, 2 ♀); Moa, Yamanigüey 75 m, 20°34'45.9"N, 74°44'10.2"W, 24/I/2009, R. Núñez, slide RNA264(wings), DNA voucher PM157-05 (2 ♂, 1 ♀); ♀), Sierra de Cristal, cerca de la Estación La Zoilita 400 m (20°37'41.7"N, 75°29'08.1"W), 15-20/II/2010, R. Núñez, DNA voucher PM07-22 (M040). Santiago de Cuba: Las Lagunas, 19°59'37"N, 75°47'50"W, 29/VI/1930 (1 ♂); Santa María, 20°05'N, 75°49'W, Julio 1940, slides RNA 177/178(androconial sclaes) (2 ♂, 1 ♀); same locality as for anterior, 18 May 1941, slide RNA180 (androconial scales) (1 ♂); same locality as for anterior, 20 Junio 1943 (1 ♂); same locality as for anterior, 29 Junio 1943 (1 ♂); Marimón, 27 Junio 1942, slide RNA179(androconial scales) (1 ♂); same locality as for anterior 28 Junio 1942, slide RNA205(wings) (1 ♀); Ote (currently Santiago de Cuba), Ciudamar, 19°58'41"N, 75°51'51"W, 22/IX/1950, S. L. de la Torre (1 ♀); Cuabitas (20°03'48"N, 75°48'05"W, 28/IV/1953, S. L. de la Torre (1 ♂); same locality as for anterior XII/1956, P. Alayo (1 ♀); Las Manuelas camino a Baire 420 m, 20°13'09"N, 76°21'52"W, 23/XI/1952, S. L. de la Torre (1 ♂); Pico Turquino 1972 m, 19°59'23.7"N, 76°50'11.9"W, 18/X/1966, I. García (1 ♀); Ote (currently Santiago de Cuba), Loma El Gato 1000 m, 20°00'33"N, 76°02'16"W, VIII/1942, Hno Crisogono (1 ♂); same locality as for anterior, 6/IX/1951, S. L. de la Torre, genitalia ♂ in glycerin (5 ♂, 5 ♀); same locality as for anterior, 17-20 June 1952, F. de Zayas & P. Alayo (1 ♂); same locality as for anterior, 20 June 1952, slide RNA187(wings)/222(legs & labial palpus) (2 ♂); same locality as for anterior, 25-26 Junio 1952, F. Zayas & P. Alayo (1 ♂); same locality as for anterior, 11/VIII/2008, E. Oliva, DNA voucher PM07-12 (M029) (1 ♂, 1 ♀); same locality and date as for anterior, E. Fonseca (1 ♀); Puerto Boniato, 28/XI/1950, S. L. de la Torre (1 ♂); same data as for anterior except 16/V/1953 (1 ♂); zona del Caney, Loma del Ermitaño 430 m, 20°02'38"N, 75°37'3"W, 13/III/1953 (1 ♂); Ote (currently Santiago de Cuba), Caney, Gran Piedra, 20°00'31"N, 75°42'31"W, Junio 1954, F. de Zayas & P. Alayo, slide RNA229(wings) (2 ♀); same locality as for anterior, 23/IV/1955, genitalia in glycerin (1 ♀); same locality as for anterior, VI/1962, P. Alayo, genitalia in glycerin (1 ♀); Juraguá próximo a Santiago de Cuba, 19°55'31"N, 75°38'28"W, 9/I/1968, S. L. de la Torre (1 ♂); alrededores Estación BIOECO Gran Piedra 1000 m, 20°00'31"N, 75°37'3"W, 16-18/XI/2005, R. Núñez (1 ♀); same data as for anterior except 8/III/2008, genitalia ♀ in glycerin (1 ♂, 1 ♀); same locality as for anterior, 14/VIII/2008, E. Oliva (1 ♂); km 19 carretera Gran Piedra, 12/III/2008, R. Núñez (1 ♂); Gran Piedra, El Olimpo, campamento forestal "Las Marianas", 13/III/2008, R. Núñez, DNA voucher PM15-07 (M053) (2 ♂). Guantánamo: Ote (currently Guantánamo), Guantánamo, 20°01'N, 75°12'W, 26/XI/1950, S. L. de la Torre & P. Alayo (1 ♀); Ote (currently Guantánamo), Baracoa, Loma La Farola, 1/V/1968, S. L. de la Torre (2 ♂, 1 ♀); Ote (currently Guantánamo), Cupeyal 730 m, 20°26'57"N, 75°03'38"W, VI/1971, I. García (2 ♂); Piedra La Vela 650 m, 20°24'45"N, 74°56'51"W, VII/2001, R. Núñez (2 ♂); Piedra La Vela, Loma El Mulo 615 m, 20°25'27"N, 74°54'32"W, VII/2001, R. Núñez (1 ♂); río Jaguaní, Vázquez Abajo 560 m, 20°25'15"N, 74°54'33"W, Cuchillas del Toa, Boca de Jaguaní 130 m, 20°22'46"N, 74°41'36"W, VIII/2001, R. Núñez (1 ♂); Yumurí del Sur 450 m, 20°11'21"N, 74°29 31"W, 20/I/2009, R. Núñez & E. Oliva (2 ♂, 2 ♀). CZACC, MFP.

Distribution.

The species is present across the Cuban archipelago from coastal areas to mountains up 1100 m (Figs 56-59).

Immature stages.

Egg & oviposition - Eggs are laid loose, near spherical in shape and ivory white in color becoming beige with irregular orange brown spots a day after laid. Torre (1968) also mentioned that eggs are laid loose. Surface is covered by a fine raised reticulation forming minute polygonal areas ( Dethier 1940, Torre 1968). Time to hatch 7 to 9 days (n=16), according Dethier (1940) 6 to 11 and Torre (1968) gave 5 to 8 days.

First instar larva (Fig. 90) - Head capsule dark brown, almost black, with a bronze gloss and with two short horns on top. Body beige, greenish white after fed on host leaves, with a dorsal line and four pairs of longitudinal pale brownish green thin lines all of same width and more or less equally spaced: subdorsal, suprastigmatal, stigmatal and infrastigmatal. Dimensions (n=16): head capsule width 0.52-0.57 mm, head capsule height 0.56-0.59 mm, initial total length 2.2-2.5 mm, final total length 3.4-3.7 mm. Duration (n=16): 7-10 days. This description agrees with that by Dethier (1940), who reported an instar duration of 7 days.

Second to fourth instars (Fig. 91) with the same pattern of fifth, described below, but paler and less contrasting.

Fifth instar larva (Figs 92, 94) - Pale morph. Head capsule pale brownish gray with numerous slightly darker dots, base of setae dark brown, a vertical brown line from each side reaching horns and almost joining at epicranial suture, horns reduced; stemmatal area, clypeus and area around mandibles brown or dark brown; mandibles amber brown, black at edge; X–mark of epicranium slightly darker than background with lower arms longer and rounded at tip, broken as four spots in some specimens. Body pale brownish yellow minutely striated in brownish gray thin lines on dorsum between subdorsal lines, with a dorsal line and five pairs of longitudinal pale brownish gray lines: subdorsal, suprastigmatal, stigmatal and infrastigmatal; dorsal line brownish gray edged at beginning of each segment by two black dots; subdorsal lines somewhat diffuse toward segments margins, with a black dot on its lower edge at posterior margin of each segment, dots on thorax enlarged, lines ending at caudal tails; suprastigmatal lines dark brown, thin, above it on each segment a central white dot encircled in black and another, black near posterior margin; stigmatal lines dark brown, thin, space between it and suprastigmatal pale beige, contrasting, edged on its lower edge by spiracles which are dark and encircled in grayish white; infrastigmatal lines thin, somewhat diffuse; subventral lines thick, wavy, and darkest; ventral side, including prolegs pale brownish yellow. Dimensions (n=4): head capsule width 1.41-1.57 mm, head capsule height 1.55-1.62 mm, initial total length 12-15 mm, final total length 20-23 mm. Duration (n=9): 11-18 days. Larvae reared by the senior author match Dethier (1940) descriptions of instars two to fourth, in general, color pattern is about the same, including the fifth instar, with minor variations.

Dark morph (Fig. 93). Head with all tones darkened. Body background pale brown with lines dark brown, somewhat diffuse; dots at edges of mid dorsal and subdorsal and encirclement of spiracles ashy white, contrasting; a thin pale yellowish beige line between subdorsal and suprastigmatal line, contrasting; dots above suprastigmatal line and encirclement of white dots above it indistinct; space between infrastigmatal and subventral offline pale yellowish beige, contrasting; subventral line thicker than in pale morph, dark brown extending over dorsum of prolegs. Torre (1968) apparently also reared larvae of this morph but only mentioned the general darkening of coloration.

Pupa (Figs 95-97) - Entirely more or less uniform stramineous; one pair of black dots at first third of legs sheaths; abdomen with a transverse ridge with a pair of more prominent crests on dorsum of segments 1 to 6; last abdominal segment short and stout, cremaster area enlarged, broad. Three days before emergence color turns brown on dorsum extending gradually to occupying entire surface. Dimensions (n=9): total length 10-11 mm, maximum width 3.5-4.5 mm. Duration (n=9): 8-10 days.

Habitat and biology.

Calisto herophile inhabits many habitats, from suburban areas at major cities to the edges of evergreen and rainforests up to 1100 m of altitude, always disturbed in some degree. Individuals can be found any month of the year throughout the island. The species is one of the commonest butterflies in Cuba, especially in altered land with predominantly herbaceous vegetation but shaded to some degree ( Fontenla 1987a; Núñez and Barro 2003; Fernández 2007). Fernández (2007) recorded it in Camagüey province from groves, hedges and open scrub land and recorded 26 plant species as nectar sources. We recorded two predation events on this species, one in November 2008 at La Chata, La Habana province, by a crab spider, Thomisidae (Fig. 98); the other in July 2009 at Pan de Matanzas, Matanzas province, by a nymph of the mantid Stagmomantis domingensis Palisot de Beauvois (Fig. 99).

Larvae eat the entire corion after hatching and feed at night, remaining in the lower parts of grasses during the day. They accepted well the substitute grasses supplied. Duration of first three instars was about one to one and half weeks each whereas the last two were around two weeks each. The prepupal stage duration was one day long and the pupal stage extended for eight to ten days. Immature development takes 60 to 70 days and goes through five larval instars. Adult emergence occurred after mid day. Dethier (1940) apparently did not complete the life cycle, describing it only to the fourth instar without mentioning the pupa or adult emergence. Dethier used several grass species as food and said that the larvae preferred lawn grass; however, he did not give scientific names of any grass species. Torre (1968), although successful in rearing the species, only described the cycle superficially and mentioning the duration, 70 to 73 days, and number of larval instars, four. He used as substitute food Saccharum officinarum , Zea mays , and Stenotaphrum secundatum , and noted that larvae grew slower with the first.

Remarks.

Calisto herophile is one of the easiest to recognize among all Cuban Calisto species. Its smaller size on average, as well as its pale wing pattern allow their unequivocal identification, although some specimens from altitudes above 800 m can be distinctly larger. The genitalia and immature stages can be also diagnostic. The species has a wide ecological range and tolerance to anthropogenic habitat alteration.

The status of Calisto herophile subspecies, Calisto herophile parsonsi Clench, 1943and Calisto herophile apollinis , is yet pending further investigation. In the present study, only old material of parsonsi was available. The unique morphological difference with the nominal subspecies is the more homogeneous pattern at UN of wings, as pointed out by Clench (1943). Genitalic comparisons revealed an identical morphology. We were able to sequence a small fragment (337 bp) of COI for two specimens of the Bahamian subspecies Calisto herophile apollinis Bates. These specimens were clearly quite different to Cuban Calisto herophile (Fig. 66) and might warrant species status. Future studies involving fresh specimens, immature stages and DNA data could clarify the status of both of these taxa.