Siderone galanthis galanthis

Dias, Fernando Maia Silva, Oliveira-Neto, José Francisco de, Casagrande, Mirna Martins & Mielke, Olaf Hermann Hendrik, 2015, External morphology of immature stages of Zaretis strigosus (Gmelin) and Siderone galanthis catarina Dottax and Pierre comb. nov., with taxonomic notes on Siderone (Lepidoptera: Nymphalidae: Charaxinae), Revista Brasileira de Entomologia 59 (4), pp. 307-319 : 315-318

Siderone galanthis galanthis ( Brazil, Maranhão)

species ( Dias et al., 2012; Dias et al., unpublished data). No different from most species of Zaretis , Z. strigosus is highly intraspecific variable, and specimens from different locations are widely different in appearance. However, this species can be easily distinguished from other South American species east of the Andes of Zaretis by the shape of the uncus, thick and strongly keeled ( Fig. 43 , C. Mielke et al., 2004). Specimens of Z. strigosus from south and southeastern Brazil also can be distinguished by the coloration of the male wings upper side ( Fig. 1 ), light orange with dull and faint brown or reddish brown markings, and of the female, uniformly light yellow, with dull and faint brown or dark brown markings ( Fig. 3 ), and the forewing underside of the female, more or less uniformly light yellow (i.e. without a clear two-color pattern), speckled with brown and dark brown ( Fig. 4 ). Although some species of Zaretis are similarly widespread and sympatric with Z. strigosus in other areas, Z. strigosus appears to be the only species of the genus occurring in southern Brazil in the coastal Ombrophilous Dense Forest and interior Ombrophilous Mixed Forest of Paraná , Santa Catarina and Rio Grande do Sul states.

Siderone galanthis catarina Dottax and Pierre, 2009 comb. nov. ( Figs. 5–8 , 30–32 , 35–36 , 40–42 , 44–57 , 61 )

Biology: similar to Z. strigosus .

Egg ( Fig. 45 ): similar to Z. strigosus in shape and color, but larger. Egg width: 1.2 mm (n = 1).

First instar: similar to Z. strigosus in shape and color, but larger. Head capsule chaetotaxy and stemmata position, and body chaetotaxy, spiracle size and relative position similar to Z. strigosus ( Figs. 23–26 ). Head capsule size: 0.91 mm (n = 1). Duration: 6 days.

Second instar ( Figs. 30 and 47 ): Head capsule similar to Z. strigosus but labrum, anteclypeus, frontoclypeus and the anterior area of the epicranium dark brown, lateral and posterior area of the epicranium light brown, including the posterior area of the horns; head horns similar to Z. strigosus but projecting dorsally at a wider angle; body color and shape similar to Z. strigosus . Head capsule size: 1.31 mm (n = 2). Duration: 6–7 days.

Third instar ( Figs. 31 , 48 and 49 ): Head capsule similar to the previous instar but somewhat rectangular in anterior view, with a different aspect ratio than Z. strigosus ; head horns about one-third the height of the head capsule; anterior area of the head capsule dark brown, and posterior area of the epicranium lighter, matching the color of the body; body mostly brown, speckled with light brown; T1–A1 supraspiracular line light brown; A2–A9 + 10 dorsal and supraspiracular line similar to Z. strigosus , but subtler and dorsally “V”-shaped in A2, while trapezoidal in Z. strigosus ; dorsal area of A2 anterior to the dorsal line and ventral to the supraspiracular line from A2–A4 black; A2 enlarged with a pair of large lateral fleshy projections; A2–A7 dorsal and subdorsal areas with oblique brown lines forming five lighter colored lozenges; each lozenge is formed by the posterior oblique line of one segment and the anterior oblique line of the next segment; lines from the posterior half of A7–A8 straight and continuing obliquely to the subdorsal area in A9 + 10; dorsally, A9 + 10 dark brown at the squared fleshy projection. Head capsule size: 1.83 mm (n = 4). Duration: 6–8 days.

Fourth instar ( Figs. 32 , 50 and 51 ): Head capsule shape and color similar to the previous instar, but larger and much larger than fourth instar of Z. strigosus ; posterior area of the epicranium lighter, matching the color of the body, and horns about half the height of the head capsule; body shape and color similar to the previous instar, but larger and with the color pattern more noticeable, in a lighter beige and light brown tinge; fleshy projections of A2 larger. Head capsule size: 2.79 mm (n = 4). Duration: 6–8 days.

Fifth instar ( Figs. 35, 36 , 52–54 ): Head capsule similar to the previous instar but rougher, with large anterior, lateral and posterior knobs; horns knobby at the tip and about two-thirds the height of the head capsule; mostly dark brown with light brown areas and a conspicuous whitish lateral line continuous with the supraspiracular line of the thorax ( Fig. 54 ); body shape similar to the previous instar, but fleshy projections of A2 greatly enlarged and anteriorly projected; body mostly dark brown; thoracic legs red; T1–A1 with a conspicuous whitish supraspiracular line; dorsal area of A2 anterior to the dorsal line black; A2–A7 dorsal and subdorsal areas with oblique brown lines forming five lozenges, the first, third, fourth and fifth are lighter brown, while the second is darker than the ground dorsal color; A7 anteriorly with black oblique subdorsal lines; dorsal area from the posterior half of A7–A8, and obliquely to the subdorsal area in A9 + 10 black; posterior end of the squared fleshy projection of A9 + 10 reddish; abdominal legs crochets arranged as a unisserial, triordinal interrupted mesoseries and anal legs arranged as a unisserial, triordinal mesal pennelipse. Head capsule size: 4.91 mm (n = 3). Duration: 13–22 days.

Pupa ( Figs. 40–42 , 55–57 ): Similar to Z. strigosus , but with a strong concave indentation with reddish brown markings in the middle of the thorax, conspicuous in ventral and dorsal views; lateral carinae strongly developed, with extensive yellowish green coloration along the mesothoracic wing cases; prothoracic spiracle reddish brown; meso and metathorax with paired dorsal yellowish green areas; metathoracic wing cases yellowish green. Abdomen similarly conical, but less compressed and wider than Z. strigosus in ventral and dorsal views; spiracles on yellowish green areas, surrounded by reddish brown irregular markings and over a small protuberance in A4–A7; cremaster similar in shape to Z. strigosus , but entirely brown, anal scar light brown. Length: 1.13 cm; height: 1.65 cm; cremaster height: 2.83 mm (n = 3). Duration: 14–22 days.

Discussion: The immature stages of S. galanthis catarina comb. nov. were described by Müller (1886) as S. ide (synonym with S. galanthis nemesis , Table 2). Müller (1886) did not note the indentation in the middle of the thorax of the pupa, and thus Muyshondt (1976) argued that the species described by Müller was another related species, probably of the genus Zaretis . Nevertheless, no other species of Siderone or Zaretis occurs at the location Müller reared S. galanthis catarina comb. nov., additionally, the descriptions and Fig. 26a and b , plate XIII, undoubtedly identifies the species as S. galanthis catarina comb. nov. Rydon (1971) illustrates the fifth instar and pupa of S. galanthis galanthis and S. galanthis nemesis (identified as S. marthesia and S. nemesis , respectively); the illustrations by Miss M. E. Fountaine do not bring up any significant difference between the two taxa. Rydon (1971), based mainly on immature stages, erected a new taxa, “Zaretidinae” to set apart Zaretis , Siderone and Coenophlebia from others Anaeini . To some extent, this arrangement follows Röber (1892), who considered Zaretis and Siderone as “intermediate” between Preponini and Anaeini . The immature stages of S. galanthis mexicana comb. nov. were described by Muyshondt (1976) as S. marthesia (which is in fact synonym with S. galanthis galanthis , Table 2), describing the fleshy squared projection of A9 + 10 as rounded, a claim that is not supported by his own figures ( Muyshondt, 1976: 162, Figs. 13–15 ), which clearly depicts the structure as described and illustrated here. Morais et al. (1996) illustrate the fifth instar and pupa of S. galanthis catarina comb. nov., identified as S. marthesia nemesis . The density of S. galanthis catarina comb. nov. in Distrito Federal, Brazil, appears to be low: larvae were found only in a little more than 2% of all Casearia sylvestris plants examined. Larvae are reported to feed on mature leaves and to behave in a similar fashion to the behavior described here. Parasitism by Chalcidoidea wasps was reported by DeVries (1987) and Morais et al. (1996). Teshirogi (2004) illustrates a fifth instar of Siderone from Peru, claiming it is S. galanthis ; however, the adult specimen used to illustrate the species is a male of S. syntyche mars Bates, 1860 ( Figs. 59 and 60 ); one year after, Teshirogi (2005) describes and illustrates again the immature stages of S. galanthis from Peru, but using the same male specimen of S.syntyche mars , a male of S. galanthis galanthis and a female of S. galanthis mexicana comb. nov. to illustrate the species reared. Given that S. galanthis is much more common than S. syntyche Hewitson, [1854] in the Neotropics and the larvae illustrated by Teshirogi (2004, 2005) are identical to the larvae described here, the identity of the species reared by him probably is S. galanthis galanthis . To the best of our knowledge, the immature stages of S. syntyche are still unknown. Janzen and Hallwachs (2015) illustrate the fourth and fifth instar and pupae of S. galanthis mexicana comb. nov., identified simply as S. galanthis . The species are illustrated by numerous photographs and a number of host plant records are provided, all in the Salicaceae ; several cases of parasitism by Tachinidae flies and Braconidae wasps are reported. The majority of the host plant records belongs to Casearia ( Beccaloni et al., 2008) , although there are records for species of Xylosma ( Janzen and Hallwachs, 2015) and Zuelania ( DeVries, 1986; Janzen and Hallwachs, 2015); C. sylvestris is the species more frequently cited as host plant for all subspecies of S. galanthis . The fact that there are no significant morphologic differences between immature stages of specimens from El Salvador, Costa Rica, Cuba, Trinidad, Peru, central and southern Brazil supports the claim that S. galanthis is a single, widespread species.

Taxonomic comments: Siderone galanthis catarina comb. nov. was described by Dottax and Pierre (2009) as a subspecies of S. nemesis , however, molecular evidence indicate that this taxon is conspecific with S. galanthis and that S. galanthis is a widespread, geographic and intraspecifically variable species ( Fig. 58 ): the mean distance of the COI fragment of 34 specimens of S. galanthis from Mexico, Costa Rica, Panamá, and from the east and west Amazon basin, the Cerrado and southern Brazil (Table 1) is 1.014% (standard error = 0.028) ( Fig.58 ). Dottax and Pierre (2009) recognized S.galanthis and its subspecies as distinct from S.nemesis and its subspecies, on the basis of the coloration of the upperside of the wings: S. galanthis , with a large red patch on the hindwing, often reaching the outer margin near the tornus, and females with a single and enlarged red or yellowish patch, while in S. nemesis , the hindwing patch is reduced; and the forewing coloration of the females are similar members of the Laboratório de Estudos em Lepidoptera Neotropical (Laboratory for Studies on Neotropical Lepidoptera ), UFPR, for additional help with the rearing; and the Conselho Nacional de Desenvolvimento Científico e Tecnológico – CNPq (National Council for Scientific and Technological Development) for the fellowship granted to the authors (FMSD 150542/2013-5, MMC 308247/2013-2, OHHM 304639/2014-1).

to that of the males, with two patches. However, both characters are subject to intraspecific variation, particularly the development of the hindwing patch, which is highly variable all through the range of the species. The patch of the forewing as a single patch varies from yellowish to red and in development, with transitional specimens to the two-patch phenotype in places such as southern Amazonas ( Colombia), Rondônia and northern Mato Grosso states ( Brazil), where two patches are discernible, connected by scattered red scales or red patches similar to the type of S. nemesis var. confluens Staudinger, 1887 . Most specimens from the Amazonian basin and the Guyanas correspond to the phenotype with large hindwing patch and single forewing patch in the female. There is no marked sexual dimorphism of the forewing upperside patches in specimens from the Antilles, South America west of the Andes and central, eastern and southern Brazil, but the genitalia of different phenotypes throughout the entire range of S. galanthis are alike ( Fig. 61 ). Therefore, the morphology and distribution data support the recognition of four subspecies ( Table 2): S. galanthis galanthis , as recognized by Lamas (2004) but including S. thebais C. Felder and R. Felder 1862 , S. nemesis var. confluens , S. nemesis f. leonora Bargmann, 1928 and S. nemesis f. exacta Bargmann, 1929 as synonyms; S. galanthis nemesis , as recognized by Lamas (2004); S. galanthis mexicana comb. nov. and S. galanthis catarina comb. nov., the latter two as diagnosed by Dottax and Pierre (2009) and here combined with S. galanthis . Nevertheless, since no specimens from the Antilles were included in the molecular analysis, further studies including Antillean specimens may indicate S.galanthis nemesis as a valid species.The molecular and morphologic analyses also indicate S. galanthis as clearly distinct from S. syntyche ( Figs. 58–62 , Table 2). Siderone syntyche ( Figs. 59–60 ) also can be morphologically distinguished from S. galanthis by the shape of the outer margin of the forewing, strongly convex; the red patch of the forewing upperside, as a single patch in both sexes, but usually absent or greatly reduced in the base of space CuA 2 –2A; the brown patches near the base of the hindwing underside, reduced and lighter in color; and the valva, generally wider, with a developed lobe in the ampulla ( Fig. 62 ).

Figures

Fig. 43. Male genitalia of Zaretis strigosus (Gmelin, [1790]), lateral. Scale bar= 0.5mm.

Figs. 1–4. Zaretis strigosus (Gmelin, [1790]) (Santa Catarina, Brazil). 1 and 2: Male, dorsal and ventral. 3 and 4: Female, dorsal and ventral. Scale bar =1 cm.

Figs. 5–8. Siderone galanthis catarina Dottax and Pierre, 2009 comb. nov. (Santa Catarina, Brazil).5 and 6: Male, dorsal and ventral. 7 and 8: Female,dorsal and ventral.Scale bar =1 cm.

Figs. 27–32. Head capsules of second to fourth instars in anterior view. 27–29: Zaretis strigosus (Gmelin, [1790]). 27: Second instar. 28: Third instar. 29: Fourth instar. 30–32: Siderone galanthis catarina Dottax and Pierre, 2009 comb. nov. 30: Second instar. 31: Third instar. 32: Fourth instar. Scale bar, Figs. 27, 30 =0.25 mm, Figs.28, 29, 31 and 32 =0.5mm.

Figs. 33–36. Head capsules of fifth instars in anterior and lateral views. 33 and 34: Zaretis strigosus (Gmelin, [1790]). 35 and 36: Siderone galanthis catarina Dottax and Pierre, 2009 comb. nov. Scale bar =1 mm.

Figs. 37–42. Pupae in ventral, dorsal and lateral views. 37–39: Zaretis strigosus (Gmelin, [1790]). 40–42:Siderone galanthis catarina Dottax and Pierre,2009 comb. nov. Scale bar= 0.5cm.

Figs. 44–57. Host plant and immature stages of Siderone galanthis catarina Dottax and Pierre,2009 comb. nov. 44: Host plant, Casearia sylvestris (Salicaceae). 45: Egg,lateral. 46: Second instar resting in frass chain. 47: Second instar, dorsal. 48 and 49: Third instar, lateral and dorsal. 50 and 51: Fourth instar, lateral and dorsal. 52 and 53: Fifth instar, lateral and dorsal. 54: Fifth instar, head capsule in anterior view. 55 and 56: Pupa, ventral, dorsal and lateral views. Scale bars= 1 mm.

Figs. 61–62. Male genitalia of species of Siderone Hübner, [1823] in lateral view. 61: Siderone galanthis catarina Dottax and Pierre, 2009 comb. nov. (Santa Catarina, Brazil). 62: Siderone syntyche mars Bates, 1860 (Huánuco, Peru). Scale bar= 1 mm.

Figs. 23–26. Schematics of the first instar of Zaretis strigosus (Gmelin, [1790]) and Siderone galanthis catarina Dottax and Pierre, 2009 comb. nov. 23 and 24: Head capsule, anterior and posterior. 25: Prothoracic plate, dorsal. 26: Thorax and abdomen, lateral.

Figs. 9–22. Immature stages of Zaretis strigosus (Gmelin, [1790).9 and 10: First instar, lateral and dorsal.11 and 12: Second instar, lateral and dorsal.13 and 14: Third instar, lateral and dorsal. 15 and 16: Fourth instar, lateral and dorsal. 17 and 18: Fifth instar, lateral and dorsal. 19: Fifth instar, head capsule, anterior view. 20–22: Pupa, ventral, dorsal and lateral views. Scale bars= 1 mm.

Figs. 59–60. Male specimen of Siderone syntyche mars Bates, 1860, dorsal and ventral (Huanuco, Peru). Scale bar= 1 cm.

Fig. 58. Neighbor joining tree using Kimura-2-Parameter distance model of 37 sequences of species of Siderone Hübner [1823] and Coenophlebia C.Felder and R.Felder,1862, with 658 base pairs of the mitochondrial gene cytochrome oxidase, Subunit I. Scale bar = 1% of distance.