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Journal of Science: Vol. 13 | Article 35 Bonfantti et al.

Male genitalia of neotropical : A comparative analysis of character variation

Dayana Bonfantti1a*, Mirna Martins Casagrande1b, Olaf Hermann Hendrik Mielke1c

1Departamento de Zoologia; Setor de Ciências Biológicas; Universidade Federal do Paraná; C. P.: 19020; 81531- 980, Curitiba, Paraná,

Abstract Charaxinae (: ) has a worldwide distribution, although it occurs mainly between the tropics. Most occur in the Neotropics, where three tribes, Preponini, Anaeomorphini, and Anaenini, can be found. Collectively, these three tribes encompass 109 species. Because of its relevance to systematics and , the male genitalia of Lepidoptera have been extensively studied. The male genitalia are composed of the last two abdominal segments and their modifications for mating, known as claspers of the bodies. In order to improve upon the systematic classification of the subfamily, 31 species of 13 genera of Neotropical Charaxinae were analyzed. All characters relevant to species and generic taxonomy were analyzed. Most structures showed morphological variations among tribes, genera, and species. These variations demonstrated to be important to Preponini, because the structural patterns of the genitalia allow the separation in two groups, Boisduval and Fruhstorfer, and are in accord with the recent taxonomic classification proposed by Ortiz- Acevado and Willmott (2013), wherein Doubleday is synonymized in Prepona and Noreppa Rydon within Archaeoprepona. In the same way, Anaeomorpha splendida Rothschild showed considerable differences from Preponini’s genera, the tribe in which it was included, confirming the revalidation of the tribe Anaeomorphini (Ortiz-Acevado and Willmott 2013). Substantial variation was found in the genital structures of , making it difficult to establish structural patterns for this group. Such structural variation, however, may be very efficient to diagnose species, such as some species of Hübner and Rydon, which can be easily identified through the presence and location of spines on the valva.

Resumo Charaxinae (Lepidoptera: Nymphalidae) tem distribuição mundial, embora ocorra principalmente entre trópicos. A maioria das espécies distribui-se na região Neotropical, onde três tribos podem ser encontradas, Preponini, Anaeomorphini, e Anaeini. Coletivamente, estas tribos abrangem 109 espécies. Devido à sua relevância para a sistemática e taxonomia, a genitália masculina de Lepidoptera, formada pelos dois últimos segmentos abdominais, tem sido extensivamente estudada. Com a finalidade de aperfeiçoar a classificação sistemática da subfamília, foram analisadas 31 espécies de 13 gêneros de Charaxinae neotropicais. A maioria das

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. estruturas apresentou variações morfológicas entre as tribos, gêneros e espécies. Estas variações demostraram ser importantes para Preponini, porque os padrões estruturais das genitálias permitiram que fossem separados em dois grupos, o primeiro Prepona Boisduval e o segundo Archaeoprepona Fruhstorfer além de estar de acordo com a recente classificação taxonômica proposta por Ortiz-Acevado and Willmott (2013), em que Agrias Doubleday esta sinonimizada em Prepona e Noreppa Rydon em Archaeoprepona. Da mesma maneira, Anaeomorpha splendida Rothschild apresentou diferenças consideráveis em relação aos gêneros de Preponini, tribo na qual estava incluída, confirmando a revalidação da tribo Aneomorphini (Ortiz-Acevado and Willmott 2013). Variações substanciais nas estruturas genitais de Anaeini foram encontradas, tornando-se difícil estabelecer padrões para esta tribo, embora tais variações estruturais possam ser muito eficientes para diagnosticar espécies, tais como algumas espécies de Memphis Hübner e Fountainea Rydon, facilmente identificadas através da presença e localização de espinhos na valva.

Keywords: Anaeini, Anaeomorphini, , morphology, Preponini Correspondence: a [email protected], b [email protected], c [email protected], *Corresponding author Editor: Carla Penz was editor of this paper. Received: 2 August 2011 Accepted: 13 March 2012 Copyright: This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits unrestricted use, provided that the paper is properly attributed. ISSN: 1536-2442 | Vol. 13, Number 35 Cite this paper as: Bonfantti D, Casagrande MM, Mielke OHH. 2013. Male genitalia of neotropical Charaxinae: A comparative analysis of character variation. Journal of Insect Science 13:35. Available online: http://www.insectscience.org/13.35

Introduction sweat (Rydon 1971). Charaxinae butterflies occur mainly between the tropics, primarily The male genitalia of butterflies has been the inhabiting the Neotropical region (Mielke et subject of several very detailed studies, as its al. 2004a). morphology can provide the basis for work of taxonomy, systematic analysis, Recently, this subfamily has undergone and phylogenetic analysis (Simonsen drastic changes in its taxonomic classification. 2006a). Some recent work on the comparative Through phylogenetic study of the tribe morphology of male genitalia has Preponini, using data from DNA sequences, examined genital muscles (Simonsen 2006a, Ortiz-Acevado and Willmott (2013) classified b) and intraspecific structural Preponini as follows: two genera were variations (Caldas 1997; Tóth 2011). synonymized, Noreppa to Archaeoprepona and Agrias to Prepona; the fifth Charaxinae (Lepidoptera: Nymphalidae) are a (monotypic), Anaeomorpha splendida, had its group of medium to large, robust butterflies. tribe status revalidated to Anaeomorphini. So, They fly fast and circulate around tree the neotropical Charaxinae contained two canopies, behaviors that make them difficult tribes, 17 genera, and 109 species (Salazar and to observe. Males and females are easily Constantine, 2001; Lamas, 2004; Salazar, attracted by bait containing fruit, 2008), and today is made up of three tribes, 15 decomposing , feces, urine, and/or

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. genera, and 109 species (Ortiz-Acevado and study, the genital parts were stored in a Willmott 2013). microvial containing glycerin, which was attached to the same pin as the exemplar. The Charaxinae are characterized by the following synapomorphies: parapatagium Drawings were made using a camera lucida sclerotized (Ehrlich 1958); forewings with R4 attached to a dissecting microscope. The main and R5 longer than their common branch; R4 lines were inked and watercolor was used to curved downward near its end, which may be mark each sclerite with a different color. at the costal margin, the apex, or the outer margin of the wing; posterior discal cell Illustrations were coded as follows: full lines closed; anepisternum II present as a distinct represent the boundaries of morphological sclerite; and mesothoracic pre-episternum structures on the first focal plane; dashed lines well-developed (Ackery 1984). These represent the boundaries of structures that lay synapomorphies were corroborated in a series underneath the main structure under study. As of contributions by Mielke et al. (2004a, b, c), with structures represented by dotted lines, who also found some variation in the way structures represented by dashed lines were vein R4 ends in itys itylus (Westwood) only shown when they were important for the and Prepona claudina annetta (Gray). interpretation of the genital anatomy. Sclerites lacking a well-defined boundary were With the goal to clarify the classification and represented by intermediate colors. Sclerites, systematics of the Neotropical Charaxinae, we and their respective color codes, are as have illustrated, compared, and ascertained follows: tegumen - yellow; saccus - cadmium variation in the male genitalia of 31 species of red; uncus - navy blue; gnathos - alizarin red; 13 genera. Some of the characters mentioned fultura inferior - turquoise; valvae - lemon have been previously mentioned in other yellow; penis - green. literature, whereas others are new. In some illustrations, the sclerites were Materials and Methods omitted in order to facilitate interpretation. For instance, the fultura inferior was omitted The species studied and material studied are from illustrations showing the lateral aspect of listed in Table 1. The specimens analyzed are the genitalia. A minimum of nine drawings retained at Coleção Entomológica Pe. Jesus were produced for each species as follows: Santiago Moure, Departamento de Zoologia, lateral, dorsal, and posterior aspects of the Universidade Federal do Paraná, and Coleção genitalia; inner and outer aspects of the valva; Prof. Dr. Olaf Mielke. and lateral (left and right), dorsal, and ventral aspects of the penis. In the descriptions and The abdomen of each exemplar was removed comparisons, only the greatest values are and immersed in potassium hydroxide10% for given for width and length. at least 24 hours (depending on the size of the abdomen) to soften the tissues. After that, the SEM photographs were taken following abdomen was washed in water. standard procedures. First, the parts were dehydrated by immersion in decreasing The genitalia were removed through a slit on ethanol dilutions (70%, 80%, 90%, and 100%) the pleura of the 7th abdominal segment. After for 10 minutes, and then immersed for an

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. additional 10 minutes in ethanol absolute. Coenophlebia archidona (Hewitson) (Figure Second, they were subjected to critical point 2 A–K). Tegumen subtriangular in lateral drying, glued to a stub, and coated. view, anterior projection wider than long, tegument fold projecting outward from The main terminology used was adopted from median to ventral region, appendices Niculescu (1978) (Figure 1), with adaptations angulares longer than wide. Arms from from Petersen (1904), Pierce (1909; 1914), tegumen and saccus distinct. Anterior Kuznetsov (1915), Snodgrass (1935), Oiticica projection from saccus elongate in lateral (1946), Okagaki et al. (1954), Sibatani et al. view; posterior projection absent. (1954), Klots (1956), Ogata et al. (1957), Snodgrass (1957), Srivastava (1965), Uncus thick, with distal portion descending, in Casagrande (1979), and Bilotta (1994). dorsal view fused with tegumen, uncus differing from tegumen by a light Results sclerotization. Anal tube developed, with ventral portion sclerotized (subscaphium). Anaeini Gnathos open, in lateral view with proximal (Fabricius) (Figure 1 A– portion entirely fused with tegumen, C- K). Tegumen triangular in lateral view, shaped, in posterior view curved with distal anterior projection and appendices angulares portion descending. wider than long. Arms from tegumen and saccus distinct. Anterior projection from Valva subrectangular, with a dorso-proximal saccus elongated in lateral view; posterior notch and a small spine on distal portion. projection shorter than anterior. Fultura inferior bilobed, V-shaped, with a deep dorsal notch. Uncus slender, with distal portion descending, in dorsal view fused with tegumen, uncus Penis: aedeagus sinuous and completely differing from tegumen by a light sclerotized, with bulbus ejaculatorius sclerotization. Anal tube reduced and entirely projecting from antero-dorsal opening. membranous. Gnathos open, in lateral view forming two distinct plates, with distal portion electra (Westwood) (Figure 3 A–K). flattened on anterior ventral portion, gnathos Tegumen subtriangular in lateral view, extended dorsoventrally toward the proximal anterior projection wider than long, tegument portion, with spines of various sizes that give fold projecting outward from anteromedian the dorsal distal section a serrated aspect. portion of arms from tegument to their apex, appendices angulares longer than wide. Arms Valva subtriangular, with inner ends from tegumen and saccus distinct. Anterior expanded. Fultura inferior forming a semi- projection from saccus elongate in lateral circle. view, about twice as long as wide; posterior projection of saccus present, shorter than Penis: aedeagus sclerotized, with bulbus anterior projection. ejaculatorius projecting from antero-dorsal opening Uncus slender, with distal portion descending, in dorsal view fused with tegumen, uncus differing from tegumen by a light

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. sclerotization. Anal tube developed, with Valva approximately rounded; inner tegument ventral portion sclerotized (subscaphium). expanded over outer tegument on ventral Gnathos open, in lateral view with the distal distal portion and bearing small spines on the portion of each gnathos close to each other, harpe. Fultura inferior bilobed. with two parts, one dorsal and one ventral distal, ventral part more developed and Penis: aedeagus completely sclerotized, with subrectangular; ventral distal part elongate bulbus ejaculatorius projecting from antero- and with a small descending apical club; in dorsal opening, posterior opening large, posterior view L-shaped aspect, expanded originating underneath the manica. apically. Fountainea glycerium cratais (Hewitson) Valva rounded, inner tegument expanded over (Figure 5 A–M). Tegumen subtriangular in outer tegument on ventral distal portion. lateral view; without anterior projection, Fultura inferior bilobed and linked ventrally appendices angulares longer than wide. Arms with the diaphragma rather than with the from tegumen and saccus distinct. Anterior valva. projection from saccus elongated in lateral view, at least three times longer than wide; Penis: aedeagus sclerotized, expanded posterior projection absent. ventrally up to insertion of manica, and bulbus ejaculatorius projecting from antero-dorsal Uncus slender, apex slender and descending, opening. in dorsal view fused with tegumen, uncus differing from tegumen by a light drurii (Butler) (Figure 4 A–K, sclerotization. Anal tube developed, with 33 C, D). Tegumen subtriangular in lateral ventral portion sclerotized (subscaphium) and view; anterior projection and appendices enveloped by the gnathos ventrally. Gnathos angulares wider than long. Arms from closed and distally forming a round plate, tegumen and saccus distinct. Anterior projecting posteriorly. projection from saccus elongated in lateral view, at least three times longer than wide; Valva subrectangular and outer tegument posterior projection present, shorter than forming a distal spine. Fultura inferior anterior projection. bilobed, with medium ventral notch, and small dorsal spines. Uncus slender, with distal portion descending, in dorsal view fused with tegumen, uncus Penis: aedeagus sclerotized, bulbus differing from tegumen by a light ejaculatorius projecting from antero-dorsal sclerotization. Anal tube developed, with opening; posterior opening irregular, with ventral portion sclerotized (subscaphium), deep notches. enveloped by the gnathos ventrally. Gnathos open, in lateral view with two parts: dorsal Fountainea halice halice (Godart) (Figure 6 and ventral-distal, both elongated, ventral- A–M). Tegumen subtriangular in lateral view, distal part descending, with a slight fold in the without anterior projection, appendices tegument. angulares longer than wide. Arms from tegumen and saccus distinct. Anterior projection from saccus elongated in lateral

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. view, at least twice longer than wide; Penis: aedeagus sclerotized, with bulbus posterior projection absent. ejaculatorius projecting from antero-dorsal opening, posterior opening sinuous. Uncus slender, with proximal ventral expansion and distal portion descending, in Fountainea ryphea phidile (Geyer) (Figure 8 dorsal view fused with tegumen, uncus A–M, 33 A, B). Tegumen subtriangular in differing from tegumen by a light lateral view, anterior projection wider than sclerotization. Anal tube developed, with long, appendices angulares longer than wide. ventral portion sclerotized (subscaphium) and Arms from tegumen and saccus distinct. enveloped by the gnathos ventrally. Gnathos Anterior projection from saccus elongated in closed, in lateral view sub rectangular, with lateral view, at least twice as long as wide; anterior and posterior margins sinuous and posterior projection absent. fused medially Uncus slender, with distal portion descending, Valva subrectangular, with small spines near in dorsal view fused with tegumen, uncus dorsal region and a distal spine. Fultura differing from tegumen by a light inferior bilobed, formed by two plates joined sclerotization. Anal tube reduced and entirely medially. membranous. Gnathos closed, in lateral view forming a single round plate, similar to the Penis: aedeagus sclerotized, with bulbus gnathos of F. glycerium cratais. ejaculatorius projecting from antero-dorsal opening. Valva subrectangular, bearing distal spine. Fultura inferior wide and semi-circular, with Fountainea nessus (Latreille) (Figure 7 A– reduced spines on apical dorsal tips. L). Tegumen subtriangular in lateral view, without anterior projection, appendices Penis: aedeagus sclerotized, with bulbus angulares longer than wide. Arms from ejaculatorius projecting from antero-dorsal tegumen and saccus fused. Anterior projection opening. from saccus elongated in lateral view, at least three times longer than wide; posterior clytemnestra forbesi Godman and projection absent. Salvin (Figure 9 A–L). Tegumen subtraprezoidal in lateral view, anterior Uncus slender, basal portion enlarged, with projection wider than long, appendices distal portion descending, in dorsal view fused angulares longer than wide. Arms from with tegumen, uncus differing from tegumen tegumen and saccus fused. Anterior projection by a light sclerotization. Anal tube developed from saccus elongated in lateral view, at least and entirely membranous. Gnathos open, in four times longer than wide, apex slender and lateral view rounded and with distal portion of curved; posterior projection absent. both gnathos close to each other, differing from the remaining species of Fountainea. Uncus thick, dorsal proximal portion crest- like, apex slender and descending, in dorsal Valva subrectangular, bearing distal spine. view not completely fused with tegumen, Fultura inferior large and bilobed. uncus with a basal fenestra, with two dorsal distal lobes. Anal tube reduced and entirely

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. membranous. Gnathos open, in lateral view Memphis acidalia victoria (H. Druce) apostrophe-like, proximal portion underneath (Figure 11 A–L). Tegumen subtriangular in tegumen, outer distal portion descending and lateral view, without anterior projection, curved, ventral portion elongated, descending, appendices angulares longer than wide. Arms with distal portions of both gnathos close to from tegumen and saccus fused. Anterior each other and enveloped in membranes. projection from saccus elongated in lateral view, distal portion enlarged; posterior Valva approximately semi-circular. Fultura projection absent. inferior reduced, V-shaped. Uncus slender, ventral margin sinuous, apex Penis: aedeagus sclerotized and sinuous, with slender and descending, in dorsal view fused bulbus ejaculatorius projecting from antero- with tegumen. Anal tube developed and with dorsal opening. ventral portion sclerotized (subscaphium). Gnathos open, in lateral view elongated, with Hypna clytemnestra huebneri Butler (Figure proximal portion wider than long, descending 10 A–L). Tegumen, subtraprezoidal in lateral distally and with tegument fold projecting view, anterior projection wider than long, outward like a shell, in ventral view with appendices angulares longer than wide. Arms conspicuous suture indicating the fusion from tegumen and saccus distinct, arms from between lateral arms of gnathos. saccus projected dorsally beyond ventral margin of tegumen. Anterior projection from Valva semi-circular, with reduced spines on saccus elongated in lateral view, four times sacculus, a small notch on dorsal margin and a longer than wide and descending; posterior spine on distal portion. Fultura inferior projection absent. bilobed.

Uncus thick, proximal dorsal portion crest- Penis: aedeagus sclerotized, with bulbus like, apex slender and descending, in dorsal ejaculatorius projecting from antero-dorsal view not completely fused, uncus with a basal opening, anterior portion curved descending, fenestra. Anal tube reduced and entirely and posterior portion with sinuous of different membranous. Gnathos open, in lateral view shapes and dimensions. apostrophe-like, proximal portion underneath tegumen, outer distal portion descending and Memphis glauce glauce (C. Felder and R. curved, elongate in ventral or posterior views, Felder) (Figure 12 A–K). Tegumen descending, with distal portion of both subtriangular in lateral view, anterior gnathos close to each other and enveloped in projection wider than long, tegument fold membranes, membranes joined distally. extended over entire anterior margin, projecting outward, appendices angulares Valva approximately rounded. Fultura inferior wider than long. Arms from tegumen and reduced and V-shaped. saccus distinct. Anterior projection from saccus triangular in lateral view, descending, Penis: aedeagus sclerotized, sinuous, with slender distally, projecting toward the left bulbus ejaculatorius projecting from antero- side; posterior projection present, shorter than dorsal opening, posterior opening sinuous. anterior projection.

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. Uncus slender, with ventral margin sinuous, areas with reduced spines on costa and apex descending, in dorsal view fused with sacculus. Fultura inferior wide and bilobed. tegumen, uncus differing from tegumen by a light sclerotization. Anal tube developed and Penis: aedeagus sclerotized, with bulbus with ventral portion slcerotized ejaculatorius projecting from antero-dorsal (subscaphium). Gnathos open, in lateral view opening. elongated, proximal portion expanded and distal portion semi-triangular, with median Memphis lemnos (H. Druce) (Figure 14 A– constriction, giving the structure a shovel-like L). Tegumen semi-trapezoidal in lateral view, appearance. without anterior projection, tegument fold projecting outward, following anterior margin, Valva subrectangular, each bearing a distal appendices angulares longer than wide. Arms spine; spines reduced on sacculus and costa. from tegumen and saccus fused. Anterior Fultura inferior with narrow stripe. projection from saccus subtriangular in lateral view, slightly longer than wide; posterior Penis: aedeagus sclerotized, with bulbus projection present, shorter than anterior. ejaculatorius projecting from anterio-dorsal opening, posterior portion with tegument fold Uncus slender, ventral margin sinuous, apex near posterior opening. slender and descending, in dorsal view fused with tegumen, but with proximal lateral Memphis hirta (Weymer) (Figure 13 A–K). portions distinct. Anal tube developed, with Tegumen subtriangular in lateral view, ventral portion sclerotized (subscaphium). anterior projection wider than long, Gnathos closed, in lateral view elongate, appendices angulares longer than wide. Arms proximal portion wider than distal; distal from tegumen and saccus distinct, arm from portion with posterior and anterior projections tegumen with small distal expansions in posterior view. projecting anteriorly. Anterior projection from saccus subrectangular in lateral view, twice as Valva subrectangular, bearing distal spine and long as wide; posterior projection present, tufts of setae on inner tegument. Fultura shorter than anterior projection. inferior bilobed.

Uncus slender, with distal portion descending, Penis: aedeagus sclerotized, with bulbus in dorsal view fused with tegumen, uncus ejaculatorius projecting from antero-dorsal differing from tegumen by a light opening, posterior-dorsal opening extended to esclerotization. Anal tube developed and with manica. ventral portion sclerotized (subscaphium). Gnathos open, in lateral view elongated, Memphis moruus stheno (Prittwitz) (Figure proximal portion wider than distal, from 15 A–K). Tegumen subtriangular in lateral posterior view with tegument fold projecting view, anterior projection wider than long, posteriorly on distal portion, shell-like. appendices angulares longer than wide. Arms from tegumen and saccus distinct. Anterior Valva semi-circular, each bearing a distal projection from saccus elongated in lateral spine, conspicuous spine on dorsal margin and view, at least three times longer than wide; posterior projection absent.

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. Penis: aedeagus sclerotized, with bulbus Uncus slender, ventral margin sinuous and ejaculatorius projecting from antero-dorsal distal portion descending, in dorsal view fused opening, posterior opening sinuous. with tegumen, uncus differing from tegumen by a light sclerotization. Anal tube developed, Memphis polyxo (H. Druce) (Figure 17 A– with ventral portion sclerotized K). Tegumen subtrapezoidal in lateral view, (subscaphium). Gnathos closed, in lateral anterior projection wider than long, tegument view elongate, proximal portion wider than fold on anterior margin, projected outward, distal, and with a tegument fold projecting appendices angulares longer than wide. Arms outward. from tegumen and saccus distinct. Anterior projection from saccus elongated in lateral Valva subrectangular, with small spines on view, twice as long as wide; posterior sacculus. Fultura inferior bilobed. projection absent.

Penis: aedeagus sclerotized, with bulbus Uncus slender, with proximal portion enlarged ejaculatorius projecting from antero-dorsal and distal portion descending, in dorsal view opening, posterior-dorsal opening extended to fused with tegumen, uncus differing from manica. tegumen by a light sclerotization. Anal tube developed, with ventral portion sclerotized Memphis philumena corita (Fruhstorfer) (subscaphium). Gnathos closed, in lateral (Figure 16 A–K). Tegumen subtrapezoidal in view elongate, proximal portion about twice lateral view, anterior projection wider than wider than distal portion, tegument fold long, tegument fold projecting outward, projecting outward. appendices angulares longer than wide. Arms from tegumen and saccus distinct. Anterior Valva subrectangular, each bearing a distal projection from saccus elongated in lateral spine, spines on sacculus and inner tegument view, about four times longer than wide; small, near costa. Fultura inferior bilobed. posterior projection present, shorter than anterior. Penis: aedeagus sclerotized and sinuous, with bulbus ejaculatorius projecting from antero- Uncus slender, with distal portion descending, dorsal opening, posterior portion with in dorsal view fused with tegumen, uncus tegument fold, posterior opening sinuous. differing from tegumen by a light sclarotization. Anal tube reduced and entirely Prozikania suprema (Schaus) (Figure 18 A– membranous. Gnathos open, in lateral view L). Tegumen subtriangular in lateral view, elongate, with proximal portion wider than without anterior projection, tegument fold distal; ventral proximal portion underneath projected outwards on anterior margin, tegumen, with distal tegument fold and dorsal appendices angulares longer than wide. Arms distal notch. from tegumen and saccus fused. Anterior projection from saccus elongated in lateral Valva subrectangular and unmodified. Fultura view, at least twice longer than wide; inferior bilobed. posterior projection absent.

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. Uncus slender, apex slender and descending, species analyzed. Fultura inferior small and in dorsal view fused with tegumen, uncus semi-circular. differing from tegumen by a light sclerotization. Anal tube developed, with Penis: aedeagus sclerotized, with bulbus ventral portion sclerotized (subscaphium). ejaculatorius projecting from antero-dorsal Gnathos closed, in lateral view elongate, opening, posterior opening sinuous. proximal portion twice as wide as distal, outer margin sinuous, distal tegument fold nemesis catarina Dottax and projecting outward. Pierre (Figure 20 A–K). Tegumen subtriangular in lateral view, anterior Valva subrectangular, distally with outer projection wider than long, appendices tegument expanded. Fultura inferior wide and angulares longer than wide. Arms from forming a semi-circle. tegumen and saccus fused. Anterior projection from saccus elongated in lateral view, at least Penis: aedeagus sclerotized, with bulbus four times longer than wide; posterior ejaculatorius projecting from antero-dorsal projection absent. opening, posterior opening irregular. Uncus slender, with distal portion descending, Pseudocharaxes xenocrates punctimarginale in dorsal view fused with tegumen, with (Kaye) (Figure 19 A–K). Tegumen triangular fenestra medially. Anal tube subtriangular in lateral view, anterior developed, with ventral portion sclerotized projection wider than long, tegument fold of (subscaphium). Gnathos open, in lateral view anterior margin projected outward, appendices elongate, completely enveloped by angulares longer than wide. Arms from membranes. tegumen and saccus separated, arms from tegumen long, the longest among the species Valva rectangular. Fultura inferior narrow and studied in this work. Anterior projection fom semi-circular. saccus semi-triangular in lateral view, about twice as long as wide; posterior projection Penis: aedeagus sclerotized, with bulbus absent. ejaculatorius projecting from antero-dorsal opening, posterior opening large and sinuous. Uncus slender, ventral margin sinuous, apex slender and descending, in dorsal view fused Zaretis isidora (Cramer) (Figure 21 A–K). with tegument, with triangular fenestra Tegumen subtriangular in lateral view, medially. Anal tube developed, with ventral anterior projection wider than long, portion sclerotized (subscaphium). Gnathos appendices angulares as wide as long. Arms closed, in lateral view elongate, with distal from tegumen and saccus distinct. Anterior tegument fold projected outward, giving the projection from saccus elongated in lateral ghathos a shell-like appearance. view, at least three times longer than wide; posterior projection from saccus absent. Valva rectangular, dorsal margin sinuous, spine on distal portion and setae on inner Uncus slender, apex slender and descending, region, setae more numerous than in other in dorsal view fused with tegumen, distinct from tegumen by light sclerotization. Anal

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. tube developed, with ventral portion opening. This is the only species studied with sclerotized (subscaphium). Gnathos open, in small spines on dorsal portion, near the source lateral view elongate, distal portion of both of the posterior opening. gnathos close to each other, tegument fold on distal portion shell-like. Preponini Archaeoprepona amphimachus pseudo- Valva rectangular, with inner and outer ends meander (Fruhstorfer) (Figure 24 A–L, 33 conspicuous. Fultura inferior narrow and E). Tegumen rectangular, anterior projection semi-circular. as wide as long, anterior tegument fold projecting outward; appendices angulares Penis: aedeagus sclerotized, with bulbus wider than long. Arms from tegumen and ejaculatorius projecting from antero-dorsal saccus fused. Anterior projection from saccus opening. subrectangular in lateral view; posterior projection present, shorter than anterior. Anaeomorphini Anaeomorpha splendida Rothschild (Figure Uncus thick, with short setae on ventral distal 22 A–K). Tegumen triangular in lateral view, portion, apex slender, in dorsal view fused anterior projection as wide as long, slightly with tegumen, uncus differing from tegumen concave on distal portion; appendices by a light sclerotization. Anal tube reduced angulares wider than long. Arms from and entirely membranous. Gnathos open, in tegumen and saccus fused. Anterior projection lateral view long, with descending projection, from saccus subtriangular, in lateral view, apex shaped like a club, concave, striated. wider than long; posterior projection present, subequal to anterior projection, with sharp tip. Valva semi-triangular, distal expansion of inner tegument with short setae, conspicuous Uncus thick, as large as tegumen, apex slender spine arising from outer tegument. Fultura and descending, in dorsal view fused with inferior bilobed with dorsal median notch. tegumen, uncus differing from tegumen by a light sclerotization, setae on ventral margin at Penis: aedeagus membranous ventrally, with the beginning of tegument fold, which bulbus ejaculatorius projecting from antero- projects latero-ventrally and has a row of drosal opening. spines near distal portion. Anal tube developed and entirely membranous. Gnathos Archaeoprepona chromus chromus open, in lateral view with proximal and distal (Guérin–Méneville) (Figure 24 A–K). portions covered by the tegumen, in posterior Tegumen rectangular in lateral view, anterior view L-shaped, distal portion longer than projection wider than long, fold from anterior proximal. tegument projecting outward, appendices angulares longer than wide. Arms from Valva subtriangular, sharp on distal portion. tegumen and saccus distinct. Anterior Fultura inferior rectangular, twice as long as projection from saccus elongated dorso- tall, with concave dorsal margin. ventrally in lateral view; posterior projection shorter than anterior projection, with sharp Penis: aedeagus sclerotized, with bulbus apex. ejaculatorius projecting from antero-dorsal

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. Uncus thick, in dorsal view fused with narrow in dorsal or ventral views, bulbus tegumen, uncus differing from tegumen by a ejaculatorius projecting from antero-dorsal light sclerotization, dorsal lobe with opening. conspicuous boundaries. Anal tube reduced and entirely membranous. Gnathos open, in Archaeoprepona demophoon andicola lateral view with proximal portion elongated, (Fruhstorfer) (Figure 26 A–G, 27 A–E). distal portion descending and striated. Tegumen rectangular in lateral view, anterior projection wider than long, tegument fold Valva semi-triangular, distal expansion of reduced and covering part of the hollow, tegument, and distal spine. Fultura inferior which distinguishes the arms from tegumen; bilobed, with dorsal and ventral notches. appendices angulares wider than long. Arms from tegumen and saccus distinct. Anterior Penis: aedeagus with ventral portion projection from saccus subrectangular in membranous, bulbus ejaculatorius projecting lateral view, twice as long as wide; posterior from antero-ventral opening. projection present shorter than anterior projection, slender distally and descending. muson (Fruhstorfer) (Figure 25 A–K). Tegumen Uncus thick, in dorsal view fused with rectangular in lateral view, anterior projection tegumen, uncus differing from tegumen by a as wide as long, anterior tegument fold light sclerotization, dorsal lobe with projected outward, appendices angulares conspicuous boundaries. Anal tube reduced wider than long. Arms from tegumen and and entirely membranous. Gnathos open, in saccus fused. Anterior projection from saccus lateral view comma-like, striated on distal two absent; posterior projection present, thirds. developed, subtriangular with distal portion sharp. Valva semi-triangular, distal expansion of inner tegument, spine arising from outer Uncus thick, in dorsal view fused with tegument. Fultura inferior bilobed and dorsal tegumen, uncus differing from tegumen by a inner margins rounded. light sclerotization, dorsal lobe with conspicuous boundaries and enveloped by Penis: aedeagus wider dorsoventrally when numerous spines on lateral portion. Anal tube compared with other species of the genus, reduced and entirely membranous. Gnathos anterior ventral portion rounded, small open, in lateral view comma-like, striated on anterodorsal lobe and median constriction medial distal portion. preceding manica, ventral portion membranous, bulbus ejaculatorius projecting Valva subtriangular, distal expansions of inner from antero-dorsal opening. tegument, with spine arising from outer tegument and with long setae. Fultura inferior Archaeoprepona licomedes licomedes bilobed and articulated with ventral area of (Cramer) (Figure 28 A–L). Tegumen valva. rectangular in lateral view, anterior projection wider than long, anterior ventral fold Penis: aedeagus, ventral portion membranous, projected outward, appendices angulares medial-anterior region dilated in lateral view, wider than long, ventral margin sinuous. Arms

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. from tegumen and saccus distinct. Anterior Valva semi-triangular, distal expansion arising projection from saccus subtriangular in lateral from inner tegument, and distal spine arising view; posterior projection shorter than from outer tegument. Fultura inferior bilobed. anterior, diamond-shaped. Penis: aedeagus with a concavity on ventral Uncus thick, in dorsal view fused with median portion, membranous ventrally, tegument, uncus differing from tegumen by a bulbus ejaculatorius projecting apically. light sclerotization, dorsal lobe with conspicuous boundaries. Anal tube reduced Prepona claudina annetta (Gray) and entirely membranous. Gnathos open, in (Figure 30 A–K). Tegumen semi-trapezoidal lateral view with two flattened, folded plates in lateral view, anterior projection wider than in lateral view, distal plate descending and long, deep depression on posterior ventral striated on outer portion. portion, appendices angulares wider than long. Arms from tegumen and saccus distinct. Valva subtriangular, distal expansion of inner Anterior projection from saccus oval in lateral tegument, and distal spine arising from outer view, longer than wide; posterior projection tegument. Fultura inferior bilobed, lateral present, shorter than anterior. margins sinuous, a smooth ventral-median notch and deep dorsal median notch. Uncus thick, elevated on proximal dorsal portion, with distal portion descending, in Penis: aedeagus, ventral portion entirely dorsal view fused with tegumen, uncus membranous, bulbus ejaculatorius projecting differing from tegumen by a light apically sclarotization. Anal tube reduced, entirely membranous and linked to the gnathos. Archaeoprepona meander castorina (E. Gnathos open, in lateral view elongate, with Mayr) (Figure 29 A–K). Tegumen long arm and with spiny apical club on, rectangular in lateral view, anterior projection projected dorsally. wider than long, tegument fold small and projected outward, appendices angulares Valva triangular. Fultura inferior rectangular, longer than wide. Arms from tegumen and taller than wide, dorsally with two convex saccus distinct. Anterior projection of saccus areas covered with small spines, tegument semi-triangular in lateral view, longer than fold projecting laterally. wide; posterior projection present, shorter than anterior projection. Penis: aedeagus sclerotized dorsally and with ventral sclerotization restricted to median Uncus slender, with expansion from median to region, bulbus ejaculatorius projects from distal portion, ventral margin sinuous, apex antero-ventral opening. slender and descending, in dorsal view fused with tegumen, uncus differing from tegumen laertes (Hubner) (Figure 31 by a light sclerotization. Anal tube reduced A–K). Tegumen semi-trapezoidal in lateral and entirely membranous. Gnathos open with view, anterior projection wider than long, distal portion striated. dorsal margin with small folds, appendices angulares wider than long. Arms from tegumen and saccus distinct. Anterior

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. projection from saccus oval in lateral view; Discussion posterior projection absent. Tegumen Uncus thick, apex slender and descending, in This sclerite is structurally diverse across the dorsal view fused with tegumen, conspicuous group, with differences in at least five areas. constriction between uncus and tegumen. Four shapes have been identified in lateral Anal tube reduced and entirely membranous. view: (1) semi-trapezoidal, clustering species Gnathos open, in lateral view long, with spiny of Agrias, Prepona, and Hypna, as well as club projected dorsally, and extended beyond Memphis philumena corita, Memphis lemnos, uncus. and Memphis polyxo; (2) rectangular, common to species of Archaeoprepona and Valva triangular. Fultura inferior rectangular, Noreppa; (3) triangular, clustering Anaea taller than wide, with basal lateral folds troglodyta with Anaeomorpha splendida; and projected outward. (4) subtriangular, common among species of Anaeini, and clustering species of Penis: aedeagus sclerotized, with tegument Coenophlebia, Consul, Fountainea, fold preceding manica, bulbus ejaculatorius Prozikania, Pseudocharaxes, Siderone, projecting from antero-ventral opening. Zaretis, and most species of Memphis.

Prepona pylene pylene Hewitson (Figure 32 With a few exceptions, enumerated below, the A–K). Tegumen semi-trapezoidal, anterior anterior projection of the tegumen is projection wider than long, appendices developed. It is lacking in most species of angulares wider than long. Arms from Fountainea (the anterior projection is found tegumen and saccus distinct. Anterior only in Fountainea ryphea phidile, of the four projection from saccus oval in lateral view; species studied) and a few species of Anaeini posterior projection absent. (Memphis acidalia victoria, Memphis lemnos, and Prozikania suprema). When present, the Uncus thick, with distal portion concave and anterior projection of the tegumen is often descending, in dorsal view fused with developed, longer than wide or as wide as tegumen, uncus differing from tegumen by the long. Such projection is reduced or is as wide latter through a dorsal process on the proximal as long only in Anaeomorpha splendida, portion. Anal tube reduced and entirely Archaeoprepona amphimachus membranous. Gnathos open, in lateral view pseudomeander, and Archaeoprepna long, with spiny club projecting dorsally, and demophon muson. extended beyond uncus. Other modifications in the tegumen were the Valva triangular. Fultura inferior taller than presence or absence of a fold in the tegument wide, with lateral fold projecting outward. on the anterior margin and appendices angulares. The latter are ubiquitous among the Penis: aedeagus sclerotized, bulbus species studied and vary in their dimensions, ejaculatorius projecting from antero-ventral being wider than long, as wide as long, or opening. longer than wide.

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. the anterior projection, a state present in Saccus Anaeomorpha splendida and Archaeoprepona Some differentiation between the ventral arms domophon muson. from the tegumen and the dorsal arms from the saccus was found in 23 of the 31 species Uncus analyzed, with special emphasis for P. Among the structural differences found in the xenocrates punctimarginale, with the arms of uncus, we highlight its shape in lateral view, the tegumen separated and distant from the which can be thick (three times longer than arms of the saccus. Therefore, the fusion wide) or slender (more than three times longer between these two structures has not revealed than wide). The tribe Preponini contains only a suitable character to group species or one species with a slender uncus, genera. Archaeoprepona meander meander. All other species in this tribe, as well as the species Thirty species have a saccus with an anterior Hypna and Coenophlebia (Anaeini), have a projection. The degree of development of this thick uncus. The uncus is slender in species of projection varies among species, and six Anaea, Consul, Memphis, Fountainea, shapes have been identified for it: oval, Siderone, Prozikania, Pseudocharaxes, and common among species of Agrias and Zaretis. Prepona; rectangular, a shape found only in Anaea troglodyta; subtriangular, characteristic The uncus also varies in the presence or of Anaeomorpha splendida, Archaeoprepona absence of setae on its dorsal portion. The demophoon andicola, Archaeoprepona uncus of Anaeomorpha splendida, licomedes licomedes, Pseudocharaxes Archaeoprepona amphimachus xenocrates punctimarginale, and Memphis pseudomeander, and Prepona laertes laertes lemnos; subrectangular, as in Archaeoprepona have a dorsal seta. Other differences in this amphimachus pseudomeander, structure among species are as follows: the Archaeoprepona domophon muson, and presence or absence of crests (present only in Memphis hirta; triangular, as in two subspecies of Hypna clytemnestra), Archaeoprepona meander castorina and presence or absence of spines on the dorsal Memphis glauce glauce; and elongated, portion (present in Anaeomorpha splendida, characteristic of species of Consul, Archaeoprepona amphimachus Fountainea and Hypna, as well as pseudomeander, and Prepona laertes laerte), Coenophlebia archidona, Noreppa chromus and the presence or absence of a fenestra. chromus, Prozikania suprema, Siderone When present, the fenestra may be found nemesis catarina, Zaretis isidora, Memphis either on the central portion (Pseudocharaxes acidalia victoria, Memphis moruus stheno, xenocrates punctimarginale and Siderone Memphis phlilumena corita, and Memphis nemesis catarina) or at the base of the uncus polyxo. (Hypna clytemnestra forbesi and Hypna clytemnestra huebneri). A posterior projection arising from the saccus was present in approximately 50% of the Anal tube species studied. This projection can be shorter The degree of sclerotization of the ventral than the anterior projection, a widespread portion was the only difference found in the state, or it can be subequal to or longer than anal tube. Species that have the ventral

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. portion of the anal tube sclerotized were Siderone nemesis catarina, and Zaretis considered as having a subscaphium. This isidora; and subrectangular, as in all species character can be used to separate between of Fountainea, Coenophlebia archidona, Anaeini, Anaeomorphini and Preponini. Memphis philumena corita, Memphis moruus Species of Preponini and Anaeomorphini do stheno, Memphis lemnos, Memphis glauce not have a subscaphium, whereas species of glauce, and Prozikania suprema. Anaeini do, with two exceptions, Fountainea nessus and Fountainea ryphea phidile. Besides the shape, the structure of the valvae may also vary. For instance, the presence of Gnathos an apical projection is a variable feature The gnathos has two basic shapes in the within tribes, but stable within genera. In our species studied. It may form a handle sample, the valva of all species of enveloping the anal tube, in which case it is Archaeoprepona, Fountainea, Consul, called “single” or “close,” or it may be Prozikania, Pseudocharaxes, and Memphis divided into two distinct arms, and be called (except for Memphis moruus stheno) had an “double” or “open.” In our sample, the apical projection. gnathos is single in species of Anaeini and double in species of and Anaeomorphini Beyond the characters described above, the Preponini, with some modifications among following features have been found to be species. variable: presence or absence of spines on costa (present in Fountainea halice halice, Five aspects were identified concerning the Memphis glauce glauce, Memphis hirta, shape of the gnathos: elongated, present in Memphis polyxo); presence or absence of species of Agrias and Prepona; shaped as an spines on the saccus (present in species of anvil, unique to Anaea troglodyta; covered by Memphis); presence or absence of an apical the tegument, unique to Anaeomorpha projection of the valva, presence or absence of splendida; thick, present in Archaeoprepona setae on the inner tegument of the valva and Noreppa; with base wider than distal (present only in Memphis lemnos and portion, as in Anaeini, except for Anaea Pseudocharaxes xenocrates punctimarginale). troglodyta. Regarding the modifications on These characters can be further investigated, the distal portion, they can be smooth, with and the location and shape of spines can be spiny club, or with distal margin serrated or detailed. striated and enveloped by membranes.

Valvae Fultura inferior The valvae assume various shapes: triangular, Differences were found in the fultura inferior as in Agrias and Prepona; subtriangular, in of all species. Within this structural diversity, species of Anaea, Anaeomorpha, and we were able to devise some basic aspects. Archaeoprepona; circular, common to the One is the general shape, with six variations: species of Hypna and Consul, as well as rectangular, as in all species of Agrias and Memphis acidalia victoria, Memphis hirta, Prepona; semi-circular, as in Anaeomorpha and Memhis polyxo; rectangular, as in splendida, Fountainea ryphea phidile, Pseudocharaxes xenocrates punctimarginale, Prozikania suprema, and Pseudocharaxes

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. xenocrates punctimarginale; elongated, present only in Coenophlebia archidona; V- The observation of Rydon (1971) was shaped, present among Hypna subspecies; confirmed, because the differences of the narrow, stripe-like, unique to Memphis hirta; gnathos, club-like and striated-like, were able and bilobed, present in all other species to differentiate Archaeoprepona and Prepona. analyzed. According to the results, species of Preponini can be divided into two distinct groups, Other characters identified for the fultura Prepona (including Prepona claudina inferior were useful for grouping species, such annetta) and Archaeoprepona (including as the presence of absence of tegument folds, Archaeoprepona chromus chromus), thus uniting species of Agrias and Prepona. confirming the arrangement of the genera Another such character is the presence or proposed by Ortiz-Acevedo and Willmott absence of spines and their location. (2013).

Penis The penis presents variations in the aedeagus, Among the characters observed in this study, which can be completely sclerotized or the following putative synapomorphies membranous ventrally. A ventrally support groups within Preponini: membranous aedeagus was found in the studied species of Archaeoprepona and Archaeoprepona: Aedeagus with ventral Noreppa. portion membranous and distal portion of gnathos striated; Another character that varied in the penis was the direction of the projection from the bulbus Prepona: Presence of lateral folds in the ejaculatorius. It was ventral in species of tegument of the fultura inferior, and gnathos Agrias and Prepona, and also in Noreppa with a club covered with spines. chromus chromus. In Archaeoprepona licomedes licomedes and Archaeoprepona Morphological characters can resolve meander castorina, this projection was taxonomic problems, and can still be more ventral. It was dorsal in 25 of the 31 species exploited, thus making them more accurate studied. when included in analyses such as female genitalia, color and shape of wings, and even Final considerations the shape and color of odoriferous scales in The present analysis revealed the importance males. of conducting detailed morphological analyses that focus on certain tagmas, particularly for The structural modifications of Anaeomorpha groups whose morphology has been splendida confirm a tribe, although in some neglected, such as the Charaxinae. It is aspects it resembles Anaea troglodyta, which possible that this group has been poorly deserves attention and further research about studied because specimens are rare and the degree of closeness between these two consequently they are considered precious. species belonging to different tribes, For this reason, their taxonomy has been Anaeomorphini and Anaeini, respectively. based on venation and coloration of the wings only.

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. Contrasting with the genitalia of species of Tecnológico for the stipends granted to the Preponini, the gentialia of Anaeini species is authors. variable, making it difficult to establish structural patterns within this group. However, References it is possible to distinguish among species based on some genitalic characters. For Ackery PR. 1984. Systematic and faunistic instance, species of Memphis and Fountainea studies on butterflies. In: Vane-Wright RI, are easily separated by the presence and Ackery PR, Editors. The Biology of location of spines on the valva. For Comstock Butterflies. Academic Press. (1961), the great variety of forms of gnathos, which despite having a structural pattern as a Bilotta IG. 1994. Morfologia comparada do basal line can provide many features useful abdome das espécies sulbrasileiras de for the determination of species. Morphinae (Lepidoptera, Nymphalidae). Revista Brasileira de Zoologia 11: 737–748. The results obtained herein reflect detailed dissections and observations of structures Caldas A. 1997. Notes on the male genitalia from different views (positions) that had not of the Anaea ryphea – Anaea eurypyle been previously used in the taxonomy of the complex (Nymphalidae). Journal of group. Therefore, they include structures not Lepidopterists’ Society 51: 83–90. previously considered. These characters can be used in the future in a phylogenetic Casagrande MM. 1979. Sobre Caligo beltrao analysis or when devising more consistent (Illiger). IV. Morfologia externa do adulto – diagnostic characters for species and genera. abdome (Lepidoptera, Satyridae, Brassolnae). Revista Brasileira de Biologia 39: 711–716. The detailing of such structures can be geared towards the maintenance of the status of Comstock WP. 1961. Butterflies of the Prozikania and Pseudocharaxes as genera, American Tropics. The Genus Anaea, synonymized by Lamas (2004) with Lepidoptera, Nymphalidae. American , the first as Zikania (nom. nud.), Museum of Natural History. and again used by Salazar (2008). The significant differences in the genitalia of the Ehrlich PR. 1958. The comparative species of both genera as described and morphology, phylogeny and classification of illustrated in this study corroborate the the butterflies (Lepidoptera: ). observations of Comstock (1961), Rydon University of Kansas Science Bulletin 39: (1971), and Salazar (2008), justifying the 305–370. maintenance of the genera, despite being monotypical. Klots AB. 1956. Lepidoptera. In: Tuxen SL, Editor. Taxonomist’s Glossary of Genitalia in Acknowledgments . Munksgaard.

We thank the Centro de Microscopia Kuznetsov NY. 1915. Fauna of Russia and Eletrônica da Universidade Federal do Paraná Adjacent Countries, Lepidoptera I. Petrograd. for the micro photography, and Conselho Nacional de Desenvolvimento Científico e

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. Lamas G. 2004. Nymphalidae. Charaxinae. Oiticica JF. 1946. Sobre a Morfologia do Tribe Anaeini. In: Lamas G, Editor. Checklist: Penis em Lepidoptera -Boletim do Museu Part 4A. Hesperioidea – Papilionoidea. In: Nacional. Zoologia 50: 1–79. Heppner JB, Editor. Atlas of Neotropical Lepidoptera, volume 5A. Association for Okagaki H, Sibatani A, Ogata M, Okada Y. Tropical Lepidoptera/Scientific Publishers. 1954. Male Genitalia of Lepidoptera: Morphology and Nomenclature II. Mielke CGC, Mielke OHH, Casagrande MM. Morphology Significance of Sacculus and 2004a. Estudo comparado da morfologia Furca. Annals of the Entomolgical Society of externa de itylus (Westwood) e America 48: 439–442. annetta (Gray) (Lepidoptera, Nymphalidae, Charaxinae) I. Cabeça, Ortiz-Acevedo E, Willmott KR. 2013. apêndices e região cervical. Revista Brasileira Molecular systematics of the butterflies tribe de Zoologia 21: 657–369. Preponini (Nymphalidae: Charaxinae). Systematic Entomology 38: 440–449. Mielke CGC, Mielke OHH, Casagrande MM. 2004b. Estudo comparado da morfologia Petersen W. 1904. Die Morphologie der externa de Zaretis itys itylus (Westwood) e Generationsorgane der Schmetterlinge und Agrias claudina annetta (Gray) (Lepidoptera, ihre Bedeutung für die Artbildung. Mémoires Nymphalidae, Charaxinae) II. Tórax e de L’Académie Impériale dês Sciences de St. apêndices. Revista Brasileira de Zoologia 21: Pétersbourg VIII Série XVI, 8: 1–84. 421–433. Pierce FN. 1909. The genitalia of the group Mielke CGC, Mielke OHH, Casagrande MM. of the Lepidoptera of the British 2004c. Estudo comparado da morfologia Islands. E.W. Classey. externa de Zaretis itys itylus (Westwood) e Agrias claudina annetta (Gray) (Lepidoptera, Pierce FN. 1914. The genitalia of the British Nymphalidae, Charaxinae) III. Abdome. Geometridae. Liverpool. Revista Brasileira de Zoologia 21: 905–912. Rydon AHB. 1971. The Systematics of the Niculescu EV. 1978. L’armature génitale chez Charaxidae (Lep: Nymphaloidea). les Lépidoptères. Bulletin de La Société Entomologist’s Record and Journal of Entomologique de Mulhouse (Supplement): Variation 83: 219–233, 311–316, 336–341, 1–40. 384–388.

Ogata M, Okada Y, Okagaki H, Sibatani A. Salazar JAE, Constantino LM. 2001. 1957. Male Genitalia of Lepidoptera: Synthesis of the description of new genera for Morphology and Nomenclature III. South America: Rydonia, Annagrapha, Appendages Pertaining to the Tenth Somito. Pseudocharaxes, Muyshondtia, Zikania Annals of the Entomolgical Society of (Lepidoptera, Nymphalidae). Lambillionea America 50: 237–244. (supplement 2) 3: 344–370.

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Journal of Insect Science: Vol. 13 | Article 35 Bonfantti et al. on the wing pattern and behavior of several Snodgrass RE. 1935. Principles of insect genera (Lepidoptera: Nymphaloidea, morphology. McGrow – Hill Book Company. Charaxidae). Boletin Científico Museo de Historia Natural, Universidad Caldas 12: Snodgrass RE. 1957. A revised interpretation 171–205. of the external reproductive organs of male insects. Smithsonian Miscellaneous Sibatani A, Masami O, Yoshio O, Hiromo O. Collections 135: 1–60. 1954. Male genitalia of Lepidoptera. Morphology and Nomenclature. I. Divisions Srivastava KP. 1965. Studies on lemon– of the valvae in Rhopalocera, Phalaenidae butterfly, demoleus L. (Lepidoptera) (Noctuidae) and Geometridae. Annals of the Part V – skeleton-muscular system of Entomological Society of America 47: 93– abdomen. Zoologischer Anzeiger 177: 217– 106. 236.

Simonsen TJ. 2006. Glands, muscles and Tóth JP, Vargas Z. 2011. Inter-intraspecific genitalia. Morphological and phylogenetic variation in the genitalia of the “Melitaea implications of histological characters in the phoebe group” (Lepidoptera, Nymphalidae). male genitalia of Fritillary butterflies Zoologischer Anzeiger 250: 258–268. (Lepidoptera: Nymphalidae: Argynini). Zoologica Scripta 35: 231–241.

Simonsen TJ. 2006. The male genitalia segments in fritillary butterflies: Comparative morphology with special reference to the “rectal plate” in Issoria (Lepidoptera: Nymphalidae). European Journal of Entomology 103: 425–432.

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Table 1. Species and materials studied.

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Figure 1. Anaea troglodyta (Fabricius). A. Dorsal. B. Ventral. C–K Genitalia: C. lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 2. Coenophlebia archidona (Hewitson). A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 3. Consul electra (Westwood). A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 4. Consul fabius drurii (Butler). A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 5. Fountainea glycerium cratais (Hewitson). A. Dorsal. B. Ventral. C–M Genitalia: C. Lateral view. D. Posterior view. E. Fultura inferior. F. Dorsal view. G. Gnathos. H and I Valva: H. Internal. I. External. J–M: Penis: J. Right lateral view. K. Left lateral view. L. Dorsal view. M. Ventral view. Scale bar: A and B = 1cm. C–M = 1mm. High quality figures are available online.

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Figure 6. Fountainea halice halice (Godart). A. Dorsal. B. Ventral. C–M Genitalia: C. Lateral view. D. Posterior view. E. Fultura inferior. F. Dorsal view. G. Gnathos. H and I Valva: H. Internal. I. External. J–M: Penis: J. Right lateral view. K. Left lateral view. L. Dorsal view. M. Ventral view. Scale bar: A and B = 1cm. C–M = 1mm. High quality figures are available online.

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Figure 7. Fountainea nessus (Latreille). A. Dorsal. B. Ventral. C–L Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F. Fultura inferior. G and H Valva: G. Internal. H. External. I–L Penis: I. Right lateral view. J. Left lateral view. K. Dorsal view. L. Ventral view. Scale bar: A and B = 1cm. C–L = 1mm. High quality figures are available online.

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Figure 8. Fountainea ryphea phidile (Geyer). A. Dorsal. B. Ventral. C–M Genitalia: C. Lateral view. D. Posterior view. E. Fultura inferior. F. Dorsal view. G. Gnathos. H and I Valva: H. Internal. I. External. J–M: Penis: J. Right lateral view. K. Left lateral view. L. Dorsal view. M. Ventral view. Scale bar: A and B = 1cm. C–M = 1mm. High quality figures are available online.

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Figure 9. Hypna clytemnestra forbesi Godman and Salvin. A. Dorsal. B. Ventral. C–L Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F. Fultura inferior. G and H Valva: G. Internal. H. External. I–L Penis: I. Right lateral view. J. Left lateral view. K. Dorsal view. L. Ventral view. Scale bar: A and B = 1cm. C–L = 1mm. High quality figures are available online.

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Figure 10. Hypna clytemnestra huebneri Butler. A. Dorsal. B. Ventral. C–L Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F. Fultura inferior. G and H Valva: G. Internal. H. External. I–L Penis: I. Right lateral view. J. Left lateral view. K. Dorsal view. L. Ventral view. Scale bar: A and B = 1cm. C–L = 1mm. High quality figures are available online.

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Figure 11. Memphis acidalia victoria (H. Druce). A. Dorsal. B. Ventral. C–L Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F. Fultura inferior. G and H Valva: G. Internal. H. External. I–L Penis: I. Right lateral view. J. Left lateral view. K. Dorsal view. L. Ventral view. Scale bar: A and B = 1cm. C–L = 1mm. High quality figures are available online.

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Figure 12. Memphis glauce glauce (C. Felder and R. Felder). A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 13. Memphis hirta (Weymer). A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 14. Memphis lemnos (H. Druce). A. Dorsal. B. Ventral. C–L Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F. Fultura inferior. G and H Valva: G. Internal. H. External. I–L Penis: I. Right lateral view. J. Left lateral view. K. Dorsal view. L. Ventral view. Scale bar: A and B= 1cm. C–L= 1mm. High quality figures are available online.

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Figure 15. Memphis moruus stheno (Prittwitz). A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 16. Memphis philumena corita (Fruhstorfer). A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B= 1cm. C–K= 1mm. High quality figures are available online.

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Figure 17. Memphis polyxo (H. Druce). A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B= 1cm. C–K= 1mm. High quality figures are available online.

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Figure 18. Prozikania suprema (Schaus). A. Dorsal. B. Ventral. C–L Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F. Fultura inferior. G and H Valva: G. Internal. H. External. I–L Penis: I. Right lateral view. J. Left lateral view. K. Dorsal view. L. Ventral view. Scale bar: A and B = 1cm. C–L = 1mm. High quality figures are available online.

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Figure 19. Pseudocharaxes xenocrates punctimarginale (Kaye). A. Dorsal. B. Ventral. C–K Genitalia: C. lateral view. D. posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. right lateral view. I. left lateral view. J. dorsal view. K. ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 20. Siderone nemesis catarina Dottax and Pierre. A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 21. Zaretis isidora (Cramer). A. Dorsal. B. Ventral. C–K Genitalia: C. lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 22. Anaeomorpha splendida Rothschild. A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 23. Archaeoprepona amphimachus pseudomeander (Fruhstorfer). A. Dorsal. B. Ventral. C–L Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F–H Valva: F. Internal. G. External. H. Dorsal. I–L: Penis: I. Right lateral view. J. Left lateral view. K. Dorsal view. L. Ventral view. Scale bar: A and B = 1cm. C–L = 1mm. High quality figures are available online.

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Figure 24. Archaeoprepona chromus chromus (Guérin–Méneville). A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 25. Archaeoprepona demophon muson (Fruhstorfer). A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B= 1cm. C–K= 1mm. High quality figures are available online.

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Figure 26. Archaeoprepona demophoon andicola (Fruhstorfer). A. Dorsal. B. Ventral. C–G Genitalia: C. Lateral view. D. Dorsal view. E–G Valva: E. Internal. F. External. G. Dorsal. Scale bar: A and B = 1cm. C–G = 1mm. High quality figures are available online.

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Figure 27. Archaeoprepona demophoon andicola (Fruhstorfer). A–E Genitalia: A Posterior. B–E Penis: B. Right lateral view. C. Left lateral view. D. Dorsal view. E. Ventral view. Scale bar: A–E= 1mm. High quality figures are available online.

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Figure 28. Archaeoprepona licomedes licomedes (Cramer). A. Dorsal. B. Ventral. C–L Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F–H Valva: F. Internal. G. External. H. Dorsal. I–L: Penis: I. Right lateral view. J. Left lateral view. K. Dorsal view. L. Ventral view. Scale bar: A and B = 1cm. C–L = 1mm. High quality figures are available online.

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Figure 29. Archaeoprepona meander meander (Cramer). A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F–H Valva: F. Internal. G. External. H. Dorsal. I–K: Penis: I. Right lateral view. J. Left lateral view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 30. Prepona claudina annetta (Gray). A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 31. Prepona laertes laertes (Hubner). A. Dorsal. B. Ventral. C–K Genitalia: C. lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 32. Prepona pylene pylene Hewitson. A. Dorsal. B. Ventral. C–K Genitalia: C. Lateral view. D. Posterior view. E. Dorsal view. F and G Valva: F. Internal. G. External. H–K: Penis: H. Right lateral view. I. Left lateral view. J. Dorsal view. K. Ventral view. Scale bar: A and B = 1cm. C–K = 1mm. High quality figures are available online.

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Figure 33. SEM photographs. A and B: Subscaphim of Fountainea ryphea phidile. C and D: Subscaphium of Consul fabius drurii. E. Stried gantho of Archaeoprepona amphimachus pseudomeander. High quality figures are available online.

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