Memphis leonida (Stoll, 1782)

Dias, Fernando Maia Silva, Casagrande, Mirna Martins & Mielke, Olaf Hermann Hendrik, 2012, Taxonomic notes on the leafwing butterflies Memphis leonida (Stoll) and Memphis editha (Comstock), stat. nov. (Lepidoptera: Nymphalidae: Charaxinae), Zootaxa 3343 (1), pp. 16-30 : 19-22

Memphis leonida (Stoll, 1782) ( figs 1–15 )

Papilio leonida Stoll, 1782 . pp. 203, 250, pl. 388, figs C (male dorsal), D (male ventral); described from an unknown number of specimens; Surinam.

Paphia leonidas ; Westwood, 1850. p.319; misspelling.

Paphia porphyrio Bates, 1865 . p. 340; described from an unknown number of specimens; Pará, Brazil.

Anaea porphyrio ; Röber, 1916; in Seitz, A. p. 585, pl. 119a (male dorsal).

Anaea (Memphis) leonida leonida ; Comstock, 1961. p. 144, pl. 27, fig. 5 (male dorsal and ventral; type specimen of Paphia porphyrio ), fig. 6 (female dorsal and ventral); figs 99 (male venation), 100 and 100A (female venation); fig. 219 (male genitalia).

Memphis leonida f. leonida ; D’Abrera, 1988. p. 694 (male, dorsal and ventral; female dorsal).— Pyrcz & Neild, 1996, p. 107, pl. 22, figs 945–950 (male dorsal and ventral, females dorsal and ventral).

Memphis leonida f. porphyrio ; D’Abrera, 1988. p. 694 (male dorsal).— Pyrcz & Neild, 1996, p. 107, pl. 22, fig. 944 (type male dorsal).

Memphis leonida ; Lamas, 2004, p. 227.— Carsus & Choimet, 2008. p. 20–21 (female dorsal and ventral).

Memphis leonida leonida ; Lamas, 2004, p. 227; syn.: porphyrio Bates, 1865 .

Diagnosis: Most specimens of M. leonida are distinct from most species of Memphis by their reddish brown patch in the outer margin of the upper side of both wings. Males of M. laertes (Cramer) (ranges through north of the Amazon river to the Guyanas) have a similar pattern, but the patches of M. laertes are coppery and they never reach the outer margin, additionally, the underside of the wings are reddish orange and the pattern is hardly discernible. Males of M. leonida lacking the reddish brown coloring in the outer margin of the upper side of the wings can be distinguished from M. oenomais (Boisduval) , M. basilia basilia (Stoll) , M. moruus moruus (Fabricius) and M. acidalia acidalia (Hübner) by the rufous red underside of the wings. Some specimens of M. acidalia acidalia are also red on the underside ( Comstock’s (1961: pl. 28, fig. 3 ) “iron-red phase”). However, in M. acidalia acidalia , the triangular area dorsal to the emargination of the inner margin of the forewing is distinctly marked, beige and irregular from the proximal area of the emargination to the apex. Male specimens of these species can be easily distinguished from M. leonida by the genitalia: the spine at the tip of the harpe is absent in M. oenomais , M. basilia basilia and M. moruus moruus , and the ventral part of the gnathos of M. acidalia acidalia rather small when compared with M. leonida . Females of M. leonida are easily confused with females of M. laertes , M. polycarmes , M. oenomais , M. basilia basilia , M. acidalia acidalia and M. moruus moruus . The forewing outer margin of the female of M. leonida is markedly rounded (never falcate) and both wings have a red fringe, absent in M. oenomais , M. basilia basilia , M. acidalia acidalia and M. moruus moruus . The underside pattern is similar to the pattern of the males and is clearly discernible, unlike females of M. polycarmes and M. laertes . Additionally, the underside of the wings are never coppery as in females of M. polycarmes , and never have a dark band alongside the outer margin, as in females of M. laertes . Illustrations of the females of the aforementioned species are given by Carsus & Choimet (2008) for comparison.

Redescription: ♂: ( figs 1–2, 5–6 ) Forewing average length 28 mm (25–30 mm) (n=20).

Wing shape: Forewing triangular; costal margin rounded; outer margin more or less sinuous; apex slightly falcate; inner margin straight, strongly emarginated before tornus; tornus enlarged beyond the inner margin; hindwing humeral area rounded and well–developed; costal margin mostly straight; outer margin slightly curved, with a short and pointed tail at vein M 3 and smoothly right angled at the tornus; inner margin slightly curved but not emarginated.

Wing color, upper side: Ground color of both wings dark brown; wing bases with a variable iridescent dark blue or purple suffusion; forewing often with a reddish brown marking from the tornus to R 5 –M 1 space in the length of the outer margin; forewing seldom with two small blue markings in the spaces R 4 –R 5 and R 5 –M 1; hindwing often with a reddish brown marginal and submarginal markings through the outer margin; hindwing with minute ocelli in the spaces close to the outer margin, the most conspicuous in space M 3 –CuA 1; anal fold developed, with long brown scales.

Wing color, underside: Ground color of both wings rufous red with black and white random speckles; forewing with a conspicuous lighter triangular area dorsal to the emargination of the inner margin, and a straight white line of dispersed scales from the proximal area of the emargination to the apex; hindwing with minute ocelli on the spaces near the outer margin, the most conspicuous in space M 3 –CuA 1. Specimens lacking the reddish brown coloring in the outer margin of the upper side of the wings have fainter patterns and lighter shades of red on the underside, when compared to the reddish brown phenotype.

Head: Eyes naked; labial palpi rufous red, speckled with dark brown and white scales; antennal length about half of the forewing length, segments dark brown with some ventral white scales; club slender and elongated, tip reddish.

Body: Dorsal color of thorax dark brown with random iridescent green and blue scaling; ventral surface rufous red, speckled with white scales; legs dark brown, speckled with white scales. Abdomen dorsal surface uniformly dark brown; ventral surface uniformly pale brown.

Genitalia ( figs 9–12 ): Saccus semitubular; dorsal projection sinuous and projected dorsad at a right angle. Tegumen trapezoidal, dorsally straight in lateral view; attached at a suture to the anterior part of the uncus and only slightly attached to the gnathos. Gnathos bifid and produced ventrad, arms parallel; ventral part of the gnathos fused medially, trapezoidal in ventral view. Uncus semitubular and curved ventrally, with small dorsal ridge. Valva ventrally rounded and separation between the sacullus and the harpe smooth, with a pointed projection at the end of the harpe; sacullus with minute spines; ampulla developed and rounded; costa developed anteriad, with a posterior angled projection. Aedeagus long and slender, folded in the left side and slightly bifid distally. Fultura inferior bifid and rectangular.

♀: ( figs 3–4, 7–8 ) Forewing average length 30 mm (28–34 mm). (n=20).

Wing shape: As in male, except where noted. Forewing outer margin evenly rounded; apex never falcate; emargination before tornus more developed than male; hindwing larger than male; outer margin strongly rounded and slightly crenulated, with a developed and blunt tail at vein M 3, outer and inner margins obtusely angled at the tornus.

Wing color, upper side: Ground color of both wings dull brown; wing bases with a variable iridescent blue or purple suffusion, lighter than male; forewing often with light reddish brown markings from the tornus to space R 5 –M 1; forewing with three pinkish markings in the spaces R 3 –R 4, R 4 –R 5 and R 5 –M 1; hindwing often with variable light reddish brown markings in the marginal and submarginal areas through the outer margin; occeli and anal fold as in male.

Wing color, underside: Ground color of both wings light brown with dark brown and beige random speckles; forewing pattern similar to male; hindwing with light scaling forming a band through the submarginal area along the outer margin. Both wings have a red fringe along the outer margin.

Head: As in male.

Body: Dorsal color of thorax as in male; ventral surface light brown, speckled with beige scales; legs brown, speckled with beige scales. Abdomen dorsal surface as in male; ventral surface uniformly beige.

Genitalia ( figs 13–15 ): Tergum VIII triangular, attached ventrally to the sides of the lamella postvaginalis, and dorsally to the lamella antevaginalis by a slender sclerotized loop; anal papillae round and bristled, projecting the posterior apophysis; lamella postvaginalis ventral to the anal papillae and medially projected; lamella antevaginalis asymmetrical, more or less wide, with a dorsal truncated projection; left side larger than the right and ventrally subrectangular; right side subtriangular; seminal duct near the base of the ductus bursae; ductus bursae about the same length of the copus bursae; corpus bursae oval, bearing two parallel slender signa on the left side.

Distribution ( fig. 27 ): Forest habitats in the lower and middle Amazon basin and the Guyanas; Cayenne ( French Guiana); Surinam; Amazonas and Bolívar ( Venezuela); Pará, Amazonas, Amapá, Acre, Mato Grosso and Rondônia ( Brazil). Ecological niche modeling predicts the presence of M. leonida further north along the coast of Guyana, Surinam and northeast Venezuela, northern Bolivia, eastern Colombia, and the Brazilian states of Roraima and Maranhão ( fig. 29 ) (mean AUC = 0.991). Ecological niche modeling shows that the most important environmental variables to predict the distribution of M. leonida are the temperature seasonality (50.1% contribution, 85.3% permutation importance), annual precipitation (28.6% contribution, 2.1% permutation importance) and altitude (5.1% contribution, 4.9% permutation importance). The temperature seasonality appears to have the most useful information by itself; however, altitude appears to have the most information that isn't present in the other variables.

Taxonomic comments: Stoll’s (1782) original description of M. leonida is brief and does not reveal much of the species true identity. There is no mention about the number of studied specimens; so the label “ HOLOTYPE ” anonymously attached to the specimen at RMNH (Warren et al. 2011) is inaccurate. However, this specimen probably is the one used by Stoll to describe Papilio leonida , as the specimen corresponds well with the illustration in Stoll (1782) and the RMNH collection houses a number of Cramer’s and Stoll’s type material ( de Jong 1982). This specimen is here designated lectotype to avoid incorrect identifications and further uncertainty ( ICZN 1999 Art. 74.7.) and the following label will be attached: / LECTOTYPE Papilio leonida Stoll, 1782 Dias, Casagrande & Mielke, 2012 /.

Reddish brown patches in the marginal and submarginal areas of the forewing are evident in the type specimen and the illustration of the upper side of the male provided by Stoll (1782: p. 203, pl. 388, fig. C). The specimen described as a female of M. leonida by Stoll (1782) and assumed to be a female of M. laertes by Druce (1877) is in fact a male of M. laertes (Stoll 1782: p. 203, pl. 388, figs E, F); the female of M. leonida is correctly illustrated by Comstock (1961: pl. 27, fig. 6 ) and Pyrcz & Neild (1996: pl. 22, figs 947–950). Bates (1865) described Paphia porphyrio from a large series of specimens from Pará, and stated that this species was probably the same described by Stoll (1782) as M. leonida , failing to note any other differences between these two taxa. Due to lack of material and the confusion with Stoll’s (1782) descriptions, Röber (1916) referred to M. leonida as a “dubious species”, and illustrated a specimen of M. leonida with reddish brown outer margin as Anaea porphyrio ( Röber 1916: pl. 119a). Comstock (1961) stated that Bates (1865) re-described as Paphia porphyrio the species bearing the valid name M. leonida because of the unsatisfactory illustrations and the confusion with M. laertes presented in Stoll’s (1782) figures. Therefore, some of his specimens probably also had the reddish brown markings in the outer margin, as they are quite distinctive in the type and in Stoll’s (1782) illustration. These reddish brown markings are absent in the type of Paphia porphyrio ( Comstock 1961: pl. 27, fig. 5 ; Pyrcz & Neild 1996: pl. 22, fig. 944): the specimen has a purple sheen in the outer margin of both wings instead. In fact, the word “ porphyrio ” probably is a Latin derivation from the Greek word “ porphyra ” (ορφύρα), which means “purple”. The specimen illustrated by Pyrcz & Neild (1996: pl. 22, fig. 944) and Warren et al. (2011) is here designated lectotype to avoid further misidentifications ( ICZN 1999 Art. 74.7.) and the following label will be attached: / LECTOTYPE Paphia porphyrio Bates, 1865 Dias, Casagrande & Mielke, 2012 /.

Apparently both Bates (1865) and Comstock (1961) assumed that there is variation in the presence and extent of the reddish brown patches. D’Abrera (1988), however, recognized two distinct forms of M. leonida : a specimen with reddish brown patch in the outer margin, M. leonida f. leonida , and a specimen with a purple suffusion in the outer margin, M. leonida f. porphyrio . Pyrcz & Neild (1996) apparently agreed with D’Abrera (1988), designating the purple form as M. leonida f. porphyrio . There are no significant differences between the wing shape, venation or genitalia of the reddish brown and purple specimens of M. leonida , although, as previously noted by Pyrcz & Neild (1996), purple specimens are more common in the terra firme forests of middle and upper Amazon basin, and appear to be predominant further south, while reddish brown specimens are mostly known from várzea forests of lower Amazon basin and the Guyanas ( fig. 27 ).

Type material: The Lectotype of Papilio leonida has the following labels: /Type Papilio leonida Cramer / Museum Leiden HOLOTYPE Papilio leonida Cramer / Raÿe Surinam / LECTOTYPE Papilio leonida Stoll, 1782 Dias, Casagrande & Mielke, 2012 / ( RMNH). The Lectotype of Paphia porphyrio has the following labels: / Pará, L. Amazons. H. W. Bates / ♂ / ♂ Para / Paphia porphyrio Bates ♂ / Anaea porphyrio, Bates. Godman –Salvin Coll. 1917.-2/ B.M. TYPE No. Rh.10730. Paphia porphyrio , ♂ Bates/ Syntype / Type/ Illustrated in “Butterflies of Venezuela ” A. Neild 1996/ LECTOTYPE Paphia porphyrio Bates, 1865 Dias, Casagrande & Mielke, 2012 / ( BMNH). Lectotype labels will be sent to the curators of the RMNH and BMNH collections.

Examined material: VENEZUELA: Amazonas, San Carlos de Río Negro , 1 ♂, no data ( MIZA) ; Bolívar, Gran Sabana , El Jaspe, 1 ♂, no data ( MIZA) ; PERU: Chanchamayo, La Merced, 1 ♂, no data, 29896 ( MNRJ) ; BRASIL, Amapá, Serra do Navio , 1 ♂ 23–25.VII.2007, 1 ♀ 30.VII.2007 (Mielke & Casagrande leg.), DZ 20.198*, DZ 20.226* ( DZUP) ; Amazonas, Manaus, Res. Ducke , 2 ♂ 10–13.VIII.2010 (Dias & Bonfantti leg.), DZ 20.333, DZ 20.055* ( DZUP) ; Manaus , 1 ♂, no data (ex coll May), ( MNRJ) , 1 ♀, VII.1923 (Parko leg.), 6042 ( MNRJ) ; Benjamin Constant , 1 ♀ VIII.43 (Parko leg.), DZ 19.520 ( DZUP) , 1 ♂, no data (Parko leg.), 6010 ( MNRJ) ; Borba , Rio Madeira , 2 ♀, 2.II.1943 (Parko leg.), 6045, 6043 ( MNRJ) ; 1 ♂ 6.VIII.1943, 1 ♂ 19.IV.1943 (Parko leg.), DZ 20.229, DZ 20.303 ( DZUP) , 1 ♂, V.1944 (Parko leg.), ( MNRJ) , 1 ♂, no data ( MNRJ) , 1 ♂, II.1943, 1 ♂, 19.V.1943 (Parko leg.), 5965, 5982 ( MNRJ) ; Tefé, Lago Acará , Rio Madeira , 1 ♂ 2.VIII.1943, 2 ♂, 3.VIII.1943 (Parko leg.), 6009, 5968, 5969 ( MNRJ) ; Tefé ( Ega ), 1 ♂, 29.VI.1941, 1 ♀, no data (ex coll May), 29836 ( MNRJ) ; Atalaia do Norte , Rio Itacoaí, VIII.1942 (Parko leg.), 5967 ( MNRJ) ; Manicoré , 1♂, X, (ex coll Arp), 23066 ( MNRJ) ; Acre, Marechal Thaumaturgo, Alto Juruá , 1 ♂, no data (Parko leg.), 29761 ( MNRJ) ; Pará, Santo Antônio do Tauá , 1 ♂, 1 ♀ 5.VI.2004, in copula (P. Jauffret leg.), DZ 20.049*, DZ 20.504 ( DZUP) ; Óbidos, Parintins-Gurupá , 1 ♂ 4.I.1952, 1 ♂ 22.IX.1958 (ex coll Kesselring), DZ 19.398*, DZ 20.280 ( DZUP) ; Óbidos , 1 ♀ no data, DZ ( DZUP) ; 1 ♀ II.1969 (ex coll Kesselring), DZ 20.399 ( DZUP) , 1 ♂ XII.1930 (ex coll D’Almeida), DZ 20.526 ( DZUP) , 1 ♂ no data (ex coll Gagarin), DZ 19.862 ( DZUP) ; 1 ♂ 30.VIII.1950, 1 ♀ 3.IX.1950 1 ♀ 4.IX.1950 1 ♂ 9.X.1950 (Cardoso leg.), DZ 19.864, DZ 20.413, DZ 19.996, DZ 19.367 ( DZUP) , 1 ♂ "winter" 1952 (ex coll Frey), DZ 19.965 ( DZUP) , 1 ♀ 5.VIII.1968, 1 ♂ 23.X.1968 (Gentili leg.), DZ 20.387, DZ 19.432 ( DZUP) , 1 ♀ 23.VIII.1968 (Ballini leg.), DZ 20.474 ( DZUP) , 21 ♂, no data (ex coll May), DZ 20.026 ( DZUP) , 23.049, 23.050, 23.248, 23.375, 6.036, 6.037, 6.038, 6.039, 29.673, 88.585, 88.564, 29.639, 29.643, 29.640, 29.641, 29.833, 29.696, 29.647, 29.645, 29.644 ( MNRJ); Belém, Utinga , 2 ♂, 1.II.1963, 8.II.1963 (O. Mielke leg.), ( MNRJ) , 1 ♂, X.1927 (ex coll May), 45753 ( MNRJ) ; Itaituba , Rio Tapajós , no data, 5966 ( MNRJ) ; Itaituba, Monte Cristo , 1 ♀, no data (ex coll May), 29729 ( MNRJ) ; Oriximiná , Rio Trombetas , no data ( MNRJ) ; Rio Xingu , (ex coll Arp), 23265 ( MNRJ) ; Rondônia, Vilhena , 1 ♂ 26.IX.1986, 2 ♂ 10.X.1986, 2 ♂ 15.X.1986, 1 ♂, 1 ♀ 29.X.1986, 1 ♂ 4.XI.1986, 1 ♂, 1 ♀ 13.XI.1986, 3 ♂ 20.XI.1986, 1 ♂ 27.XI.1986, 1 ♂ 4.XII.1986, 1 ♂ 18.XII.1986, (Elias leg.), DZ 19.390*, DZ 19.697, DZ 20.134, DZ 19.646, DZ 20637, DZ 19.751, DZ 20.618*, DZ 20.021, DZ 20.342, DZ 20.403, DZ 20.095, DZ 20162, DZ 19.418, DZ 20.227, DZ 19.599, DZ 19.654 ( DZUP) ; Cachoeira do Samuel , Rio Jamari , 1 ♂ II.1944 (Parko leg.), DZ 19.829* ( DZUP) , 1 ♂, no data (Parko leg), 6041 ( MNRJ) , 2 ♂, IV.1944 (Parko leg.), 5962, 5963 ( MNRJ) ; Ariquemes , 1 ♂ 19.IX.1978 (ex coll Gifford), DZ 19.758 ( DZUP) ; Porto Velho , Rio Novo , 1 ♂, 8.VII.1944, 1 ♀, 16.I.1944 (Parko leg.), 5964, 6046 ( MNRJ) ; Mato Grosso, Barra do Bugres , 31–35 Km NE, 1 ♂ 28.VI.1972 (Brown & Mielke leg.), DZ 20.458* ( DZUP) .

Figures

FIGURE 27. Distribution map for Memphis leonida (Stoll) and its phenotypes and Memphis editha (Comstock), plotted over the Brazilian biomes (IBGE 2004).

FIGURES 1–8. Habitus of Memphis leonida (Stoll). 1–4. Reddish brown specimens, Óbidos, Pará. 1. Male, dorsal. 2. Male, ventral. 3. Female, dorsal. 4. Female, ventral. 5–8. Purple specimens, Vilhena, Rondônia. 5. Male, dorsal. 6. Male, ventral. 7. Female, dorsal. 8. Female, ventral. Red fringe in the outer margin of the females are evidenced by a black square. Scale bar: 1cm.

FIGURES 9–15. Genitalia of Memphis leonida (Stoll). 9–12. Male genitalia. 9. Lateral view. 10. Gnathos and uncus, posterior view. 11. Valva, lateral view. 12. Gnathos and uncus, ventral view. 13–15. Female genitalia. 13. Lateral view, right side. 14. Lateral view, left side. 15. Ventral view. Scale bars: fig. 9 = 0.5mm; figs 10–12 = 0.25mm; figs 13–15 = 1mm.

FIGURES 28–29. Representation of the Maxent models. 28. Memphis leonida (Stoll). 29. Memphis editha (Comstock), stat. nov. Warmer colors show areas with better predicted conditions; white squares show the presence locations used in the analysis.