Calisto occulta Nunez
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: 57
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|Calisto occulta Nunez|
Calisto sp., Núñez 2009: 56
Calisto occulta is more similar to Calisto muripetens and Calisto bradleyi than other Cuban relatives. Characters separating Calisto occulta from Calisto muripetens are discussed above, at the Diagnosis section of the latter. From Calisto bradleyi , Calisto occulta differs by its darker color, its androconial patch without apical lobe, the slight red suffusion below cell at the UNFW, and its proportionally larger male and female genitalia. From the remainder Cuban species and from Bahaman ones with similar pattern, Calisto occulta can be separated by having fewer white dots at the UNHW (except Calisto bruneri ), its proportionally larger male and female genitalia, and the presence of a slight red suffusion below the cell at UNFW.The Hispaniolan Calisto confusa , Calisto hysius , Calisto obscura ,and Calisto pauli are superficially similar but are smaller,and have four white dots at the UNHW.
Male (Figs 16, 18, 29, 37) - FWL: 17-20 mm. Head: antennae dark brown, UN pale yellow at basal third and UP orange at club; eyes black, hairy, delimited by a pale yellow band; labial palpi dark brown on UN, pale yellow on UP, middle and basal segments rough. Thorax: UPFW uniform grayish brown except androconial patch, dark brown almost black. Androconial patch slightly distinct from surrounding areas, about two fifths the length of FW, approximately triangular in shape with apex rounded, anterior margin entering into cell and apex reaching M3 origin (Fig. 37). UPHW darker than FW, about the same hue of androconial patch. UNFW brown, slightly posterior to pdl (Figs 18, 29); a red patch in outer half of cell with outer margin edged by dl, patch posterior margin diagonal between anterior and posterior limits of cell; a slight red scaling below cell; dl, pdl and both stl darker than background; basal third of costa and outer edge of pdl with grayish white scales; ocellus black encircled by a pale yellow ring laying M1-M3, with two white pupils laying midway between M1- M2 and M2-M3, the posterior one more basad. UNHW brown mixed with pale yellow and grayish white scales; pdl and stl outer edged with pale yellow scales around ocellus; pdl area with three white dots at M1- M2, M2-M3 and M3-Cu1, with that on the middle greatly enlarged, dots surrounded by scattered whitish lilac scales; ocellus large, broad, laying between Cu1 and Cu2, black with a bluish white pupil at base and surrounded by a yellowish ochre ring outer edged by a ferruginous suffusion; tornal lobe slightly developed, black, innerly edge with pale yellow; legs dark brown, inner side of femora pale yellow, tibiae and tarsi white on external side. Abdomen: UP dark brown, UN pale yellowish brown. Genitalia (Fig. 45): uncus having typical bird’s beak shape with a dorsal keel and gradually tapering toward apex, arched at apical half, base protuberant and rounded, separated from tegumen by a single dorsal notch; tegumen hood shaped, dorsally flat but rounded at anterior end, approximately one half the length of uncus, lateral fold narrow, extending ventrally along vinculum; gnathos spine shaped, approximately 0.3 the length of uncus; valvae elongated with a broad base, digitiform projection of valvae stout with a very broad base and slightly concave at venter, extending toward apical third of uncus, joins to main body relatively sclerotized; saccus developed, finger–like at anterior half and flattened, slightly convex, toward venter on posterior half; aedeagus robust and slightly arched ventrad at middle, straight at basal two thirds with a strong left curve at apical third in dorsal view, moderately swollen toward basal half both in lateral and dorsal view, ventrally divided from basal third to bifid terminus, ending in a pair of ventral triangular flattened processes.
Female (Fig. 17) - FWL: 18-21 mm. Similar to male except: UP of wings uniform dark brown; UNFW with red scaling below cell more distinct than in male, below lower limit of cell. Genitalia (Fig. 53): large in proportion to body; sterigmal ring rounded and well developed, dorsal crown broad and symmetrical, ring almost entirely covered by a ventral fold slightly sclerotized; inner sterigmal loop large, sclerotized, left arched in ventral view almost reaching anterior margin of ring; ductus bursae originated at left side of sterigmal ring in ventral view, membranous; ductus seminalis arising close to origin of ductus bursae; corpus bursae greatly enlarged, broad, approximately the same length of ductus bursae, signa formed by two parallel columns of numerous transverse rows of small irregular sclerotized processes.
♂: Guantánamo, Baracoa, Monte Iberia plateau, Tetas de Julia 650 m, 20°27'47"N, 74°45'13.3"W, 20/V/2007, R. Núñez, DNA voucher PM07-10 (M017). CZACC.
7 ♂, 3 ♀: Holguín, Moa, Yamanigüey 75 m, 20°34'46.5"N, 74°45'12.2"W, 24-27/IX/2009, R. Núñez, DNA voucher PM07-23 (M041) (1 ♂, 1 ♀); same data as for anterior except: ex ova, emerged 28/I/2010, DNA voucher PM07-18 (M035) (1 ♂); same data as for anterior except: emerged 31/I/2010, DNA voucher PM07-19 (M036) (1 ♀); same data as for holotype except genitalia ♂ & ♀ in glycerin, DNA voucher PM07-04 (M004) (1 ♂, 1 ♀); Monte Iberia plateau, campamento ladera norte 600 m, 20°29'25.5"N, 74°43'51.3"W, 18/V/2007, R. Núñez, genitalia in glycerin, slide RNA165(wings) (1 ♂); Baracoa, Monte Iberia, ladera norte 385 m, 20°29'53"N, 74°43'48"W, 1/V/2011, R. Núñez (3 ♂). CZACC.
The species name derives from the Latin occultus (hidden, reserved) in reference to the cryptic nature of this species that remained hidden between its sympatric congeners until the present work.
Calisto occulta is known from a few localities from the middle part of the NSB Mountains, from the Monte Iberia plateau 14 km north to near Yamanigüey, in northeastern Cuba (Figs 56, 57). It is probable that Calisto occulta is more widespread in the NSB in areas where its habitat is preserved.
Egg & oviposition - Eggs are glued to substrate, are spherical in shape and ivory white in color becoming beige with irregular orange brown spots a day after being laid. Time to hatch 8 to 9 days (n=7).
First instar larva (Fig. 83) - Head capsule pale orange beige, with two short horns on top. Body beige, bluish white after fed on host leaves, with a dorsal line and three pairs of longitudinal pale brown lines: subdorsal, suprastigmatal, and stigmatal. Dimensions (n=7): head capsule width 0.60-0.62 mm, head capsule height 0.63-0.66 mm, initial total length 2.6-2.7 mm, final total length 3.5-3.8 mm. Duration (n=7): 13-15 days.
Second to fifth instars (Fig. 84) with the color pattern similar to that of sixth, described below, but paler and less contrasting and without the tranversal ashy gray bands.
Sixth instar larva (Figs 85-87) - Head capsule beige brown with numerous dark brown dots, a vertical dark brown line from each side reaching horns and joining at epicranial suture, a dark brown line connecting horns with subdorsal lines, horns much reduced; ventral third dark brown, almost balck, with a small rounded pale beige area at frons near clypeus; mandibles amber brown; X–mark of epicranium black with lower arms longer and rounded at tip. Body pale grayish brown, yellow from above spiracles to above prolegs, ventral side, including prolegs brown; dorsum of each segment with a darker “butterfly” like mark formed by small brown striations; lines slightly darker than background, except subdorsal which is pale yellow; each abdominal segment with a transverse ashy gray band at beginning from dorsum to suprastigmatal line; dorsal line edged at beginning of each segment by two black dots encircled in ashy gray; subdorsal lines thinner than dorsal line, wavy, closer to dorsal line at posterior margin of each segment, ending on caudal tails, with black dots on its upper edge aligned with dots of dorsal and suprastigmatal lines; suprastigmatal lines thin, diffuse, above it on each segment a central white dot encircled in brown and another, brown, near posterior margin; stigmatal lines thinner passing dorsal to spiracles encircled in ashy gray; infrastigmatal line thin and diffuse. Dimensions (n=2): head capsule width 2.41-2.57 mm, head capsule height 2.53-2.68 mm, initial total length 14-16 mm, final total length 22-23 mm. Duration (n=2): 30-35 days.
Pupa (Figs 88-89) - Head and wing sheaths pale brown with a row of black dots at wing sheaths margin; three pairs of frontal black dots: one elongated on eyes and two smaller and rounded on sheaths of legs, one at first third and other nearer to apex; wing sheaths edged on thorax by a brown line; dorsum of thorax and abdomen pale yellow with transverse rows of tiny black dots, density varies between individuals giving a darker or paler appearance to abdomen; abdomen smooth, with a brown line on sides; last abdominal segment long, stout, cremaster area reduced. Three days before emergence color turns brown on dorsum extending gradually to occupying entire surface. Dimensions (n=2): total length 11-12 mm, maximum width 4.5-4.7 mm. Duration (n=2): 18-19 days.
Habitat and biology.
The species inhabits the scrub forests (charrascales) of lowlands and rainforests up to 700 m in the NSB mountain range (Figs 60-62). At Yamanigüey scrub, it flies mostly below shrub shadow avoiding the high temperatures of insolated areas.
Larvae eat the entire corion after hatching and feed at night, remaining in the lower parts of grasses during day. Calisto occulta larvae did not accept well the two grass species supplied as substitute food and only two of seven larvae survived to pupation after undergoing six instars. Duration of first four instars was about two weeks each whereas fifth and sixth took about three and five respectively. Prepupal period was two to three days long and pupal stage extended for two and a half weeks. Immature development took up to four months. Adult emergence occurred at the beginning of the afternoon, between 14:00 and 15:00. A mated pair was observed at 3:00 pm Monte Iberia north slope in May 2011.
It isremarkable that the closest species to Calisto occulta is Calisto muripetens , an inhabitant of another mountain range. The relationship between them was discussed above. In the following paragraphs we discuss the differences with the remainder Cuban taxa.
Immature stages also support species status. The first instar of Calisto occulta has a pale orange beige head capsule which is almost black in Calisto smintheus and Calisto herophile . The longitudinal lines are fewer more spaced on sides and dorsum in Calisto smintheus and Calisto occulta than in Calisto herophile . Larvae of fifth and sixth instars of Calisto occulta have transverse ashy gray bands at beginning of each segment occupying from dorsum to suprastigmatal line, those lines are absent from Calisto herophile larvae. The subdorsal brown dots at metathorax of Calisto smintheus are absent in Calisto occulta .Pupae also show differences. Those of Calisto herophile have several pair of ridges on dorsum of abdomen and are beige, almost immaculate. In Calisto occulta , the head and thorax are pale grayish brown and the abdomen, that lacks the dorsal ridges, is beige with numerous black dots and a dark brown stripe at sides. As whole, is more spotted than the pupa of Calisto herophile but less than Calisto smintheus . Pupal head and cremaster shape are also different between species. Development time and number of larval instars also differ. The complete development took 60 to 70 days in Calisto herophile and 80 in Calisto smintheus both with five instars and 99 to 120 days and six larval instars in Calisto occulta .
The DNA analyses place all Calisto occulta (5 specimens, 2 localities) together, although the nuclear data placed a specimen of Calisto brochei within the Calisto occulta clade (Fig. 66). Both datasets suggest that Calisto occulta is related to Calisto muripetens , Calisto bradleyi and Calisto herophile , perhaps with Calisto muripetens being the closest relative. The species is separated from Calisto herophile and Calisto bradleyi with an average COI distance of 2.28% and 3.09% respectively, while the average COI divergence within Calisto occulta sampled from two distinct localities is just 0.98%.
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