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Journal of the Lepidopterists' Society 42(4), 1988, 269-275
SYSTEMATIC STATUS AND DISTRIBUTION OF THE
LITTLE-KNOWN CHARAXINE PREPONA WERNERI
HERING & HOPP
Kurt Johnson
Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024
AND
Henri Descimon
Laboratoire de Systematique evolutive, Universite de Provence, 3 Place Victor Hugo, F-13331, Marseille, Cedex 3, France
ABSTRACT. Prepona werneri, hitherto of uncertain systematic status, and since 1925 recorded from only the holotype male, is authenticated from eight additional specimens. Genitalia dissection and review of characters defining Archaeoprepona Fruhstorfer and Prepona Boisduval indicates werneri belongs in Prepona sensu stricto. Most specimens are from hydric forest habitat in the Choco and Cauca areas of endemism, Colombia, but one has data indicating occurrence southward in the upper Rio Putumayo region. The disjunct distribution is probably relict, reflecting former wider occurrence of per-humid biomes.
Additional key words: Apaturidae, Archaeoprepona, Neotropics, biogeography.
Of all "Prepona" butterflies, P. werneri Hering & Hopp (1925) has been the most problematical. Previously recorded only from the holotype male (Hering & Hopp 1925, Le Moult 1932-33), its melanic appearance, unusual under-surface wing pattern, and hitherto unexamined genitalia have made it a taxon of uncertain status. The most recent treatment of Neotropical Nymphalidae (D'Abrera 1987) does not mention the species. From fieldwork and survey of public and private collections, we recently located eight additional specimens of P. werneri. Only two of these were collected since 1929, and it appears unlikely that more specimens will soon be available for study. We therefore summarize below our current determinations of the taxonomic status, biology, and biogeography of this seldom-collected charaxine butterfly.
Taxonomy of "Prepona" Butterflies
Despite accumulation of specimens in private and public collections, there has not been wide agreement on the systematics of "Prepona" butterflies. Because of overall similarity in the striking blue and black markings of the wing upper surfaces, many authors have treated "Prepona" as a monophyletic group (Comstock 1944, Barcant 1970, Brown & Heinemann 1972, Riley 1975). However, as early as 1915, Fruhstorfer defined two subgroups of "Prepona". One he described as genus Ar-
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chaeoprepona (type species demophon Linnaeus) (Fig. 2C), which he regarded as "primitive" (Fruhstorfer 1915). The other, including taxa placed with Boisduval's (1836) Prepona (type species demodice Bois-duval) (Fig. 2D), he noted as sharing all morphological characters with Agrias Doubleday, from which it differed only in wing pattern. Le Moult (1932-33) also noted the heterogeneity of the group and proposed Pseudoprepona (type species demophon L., a junior objective synonym of Archaeoprepona). The above distinctions were followed by a number of authors (Orfila 1950, Rydon 1971, Descimon et al. 1973-74, Johnson & Quinter 1982).
As defined hitherto in the literature, some obvious phenetic differences separate Ar chaeoprepona from Prepona (Table 1). Authors recognizing these differences have considered both groups as genera (Sti-chel 1939, Papworth 1982) and even tribes (Rydon 1971) (Table 1). Our concern when considering the taxonomic status of werneri has been whether Prepona and Ar chaeoprepona are monophyletic groups. Our unpublished numerical cladistic analyses on taxa placed in these groups (Table 1), including outgroups Char axes, Polyura, Palla, Eu-xanthe and Comstock's (1961) Anaea sensu lato, do not conflict with generic usage of Prepona and Ar chaeoprepona as reviewed in Table 1. Therefore, based on male genitalia (Fig. 2A, B) and historical usage, P. werneri can be reliably placed in Prepona sensu stricto.
Prepona werneri Hering & Hopp (Figs. 1A, B, C, 2A)
Diagnosis. Male. Upper surface of wings: ground darker black-brown than congeners, with blue stripes of deeper azure color (not silverish or blue-green) restricted thinly caudad the forewing discal cell and in a median arc across hindwing. Under surface of wings: hindwing with variably suffused median band, area basad discal band variously marked with dark blotches, two large eyespots each submarginal in cells RS and CuAl (or a third in cell M3), forewing with disjunct or continuous apical and postmedian lines. Genitalia (Fig. 2A). Typical of general configuration of Prepona sensu stricto (Fig. 2D).
Female. Unknown.
Distribution. Principally Choco and Cauca regions (region names follow areas of ende-mism postulated by Brown 1976, 1982), Colombia, with a single specimen having data indicating upper Putumayo region.
Known specimens. In addition to the type male (Zoologisches Museum der Humbolt Universitat zu Berlin, ZMH), labelled "Origin, Prepona werneri Hering & Hopp, Rio Micay, Columbien, Februar 1925, 1000m" (Fig. 1A, B, C), eight male specimens are reported here for the first time: (1) Rio Guayabal, Colombia, February 1929, anonymous private collection (examined by first author); (2) Rio Bravo, Prov. Valle, Colombia, March 1985, anonymous private collection (noted by collector as only specimen taken at locality in many years of collecting, examined by David Matusik, Field Museum of Natural History, FMNH, photograph examined by us); (3) Frontino, Colombia, no other data, anonymous private collection (photograph furnished to first author), one male; (4) Cis-neros, Colombia, 6 May 1928 (purchased from Le Moult collection February 1968 for Niedhoffer collection), Milwaukee Public Museum (MPM) (photograph examined; genitalia dissected, illustrated in Fig. 2D); (5) Rio Micay, Cordillera Occidentale, Colombia,
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Table 1. Main characters in literature differentiating Archaeoprepona and Prepona.
Character location (authors)
Prepona
Archaeoprepona
Wing upper surface (1-6)
Hindwing under surface (1-6)
Male genitalia (1, 3-5)
Female genitalia (3, 7)
Androconia well-defined, brushlike, with rigid setae
Eyespots large, usually two, post-median to marginal, cells RS and CuAl
Slender in all parts (especially uncus and valvae); gnathos rodlike, with prominent radial spines
Sterigma Y-shaped
Androconia diffuse, softly hairy
Eyespots small, undifferentiated, marginal, cells RS to CuA2
Stout in all parts; gnathos flat, without spines
Sterigma circular
Taxa included: Prepona amesia Fruhstorfer, brooksiana Godman & Salvin, deiphile Godart, demodice Godart, dex-amenes Herbst, eugenes Bates, garleppiana Staudinger, gnorima Bates, laertes Hiibner, omphale Hubner, pheridamas Cramer, praeneste Hewitson, pylene Hewitson, neoterpe Hewitson, xenagoras Hewitson, Archaeoprepona amphimachus Fabricius, Camilla Godman & Salvin, chalciope Hubner, demophon Linnaeus, demophoon Hubner, licomedes Cramer, phaedra Godman & Salvin, meander Cramer (Rydon 1971 included chromus Guerin-Meneville and priene Hewitson in his genus Noreppa and treated genera as tribes).
Authors: (1) Fruhstorfer (1915, 1916)***; (2) Stichel (1939)**; (3) Orfila (1950)***; (4) Rydon (1971)***; (5) Descimon et al. (1973-74)***; (6) Papworth (1982)**; (7) Johnson and Quinter (1982)*. * Emphasized certain characters, ** grouped taxa based on these characters.
February-April 1928, collector Kruger, sold by Niepelt 31 May 1928, in Biedermann Collection, Zurich, Switzerland (examined by second author); (6) Cisneros, Rio Dagua valley, 1000 m, 28 February 1928, collector Hopp, sold by Staudinger 15 May 1928, in Biedermann Collection (examined by second author); (7) Queremal, Colombia, November 1986, collector Julian Salazar, Manizales Museum (K. S. Brown Jr. pers. comm., sole South American deposition known to him); (8) Upper Rio Putumayo valley, 1981, local collectors, obtained by David Matusik (FMNH), deposited in American Museum of Natural History (AMNH) (Fig. ID).
No specimens were located at Allyn Museum of Entomology (AME), British Museum (Natural History) (BMNH), Carnegie Museum of Natural History (CMNH), Field Museum of Natural History, Rijkmuseum van Natuurlijke Historie (Leiden, Netherlands) (RMNH), Museum National d'Histoire Naturelle (Paris), National Museum of Natural History (Smithsonian Institution).
Variation. Variation in the Choco and Cauca samples appears slight (Fig. 1C), but the single Putumayo specimen (Fig. ID) is distinctive, as follows: hindwing with emphatic medial band, area basad discal band with heavy blotched markings, three large submar-ginal eyespots (cells RS, M3, CuAl), and forewing with subapical stripe connected to postmedian stripe across vein M3.
Biology. The few acquirers of P. werneri provide the only sources of information about the butterfly's biology. Most specimens now in public (6) or private (3) collections derive from the pre-World War II era of highly financed butterfly sampling in the Neotropics. Initially, commercial interest prompted collection of P. werneri at several localities on the Pacific slopes of the Colombian Cordillera (mostly Choco region). These sites proved extremely inhospitable (Hering & Hopp 1925), being rain forest with extraordinarily high precipitation; Gentry (1982) cites Choco as the rainiest tropical forest in the world. Consequently, commercial interest in the insect waned, and only one specimen has since been recorded from the region (specimen 6 above). Specimens are so few that most private owners, to avoid deluges of buy offers, request anonymity.
Biogeography. Most specimens of P. werneri are from the Choco region, though one (Queremal, Colombia) is near its eastern margin with the Cauca region. Very likely the extremely hydric Choco region was a "forest refugium" during Pleistocene glaciations
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Fig. 1. Prepona werneri. A, B, Le Moult's (1932-33) figure of the holotype of P. werneri. A, Under surface. B, Upper surface. C, Reproduction of Hering and Hopp's (1925) original figure of holotype (showing distinctive markings characterizing Choco and Cauca specimens). D, Drawing indicating distinctive markings on divergent specimen from upper Rio Putumayo region.
(Brown 1976, 1982), explaining the occurrence of a number of highly insular and seldom collected butterflies in it and immediately adjacent areas. Brown emphasizes the close geographic proximity of the Choco and Cauca regions, and includes them in his "Northwestern Region" cluster (Brown 1976). He notes zones of hybridization between their taxa. If represented only by Choco and Cauca specimens, P. werneri might be characterized as a seldom collected, insular cis-Andean species, typifying limited hydric habitat. However, a larger view of its taxonomy and biogeography is necessitated by specimen 8 above from the upper Putumayo region of south-central Colombia. This collection is
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Fig. 2. Male genitalia of Archaeoprepona and Prepona, and male genitalia and abdominal androconia of P. werneri. A, Topotypical P. werneri, lateral view of genitalia with aedeagus removed (aedeagus, lateral view, beneath) and (x) ventral view, juxta, (y) lateral view, abdominal androconia at first and second abdominal spiracles. B, P. werneri specimen from upper Bio Putumayo region (dashed lines indicating areas of genitalia not available for study because of prior damage to abdomen). C, Archaeoprepona, type species demophon, Bio de Janeiro, Brazil, same format except for x and y. D, Prepona type species demodice, Bio de Janeiro, Brazil, same format except for x and y. Females of Archaeoprepona and Prepona are illustrated in Orfila (1950).
particularly striking, since the Andes are usually considered as a very efficient barrier against faunal exchange. The Putumayo region is located disjunctly southwest of the Choco and Cauca regions and included in Brown's (1976) "Andean Foothills" cluster. Brown notes very little hybridization between taxa of the Putumayo and Choco-Cauca regions. Faunal elements of the Putumayo region are mostly trans-Andean. Thus, occurrence of P. werneri in the Putumayo region appears biogeographically significant. It seems likely that disjunct distribution in P. werneri is relict, reflecting former more widespread occurrence of perhumid biomes. Compared to the rest of the Andes, uplift of its northern elements was relatively recent (Gansser 1973). Consequent separation of P. werneri into cis-Andean and trans-Andean nuclei associated with general climatic drying appears more likely than dispersal across the Andes in present or recent times. If further documented, the Putumayo P. werneri could be construed as a subspecies.
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Acknowledgments
Dale Jenkins (AME), K. S. Brown Jr. (Universidade Estadual de Campinas, Brazil) and two anonymous reviewers made helpful comments on the manuscript. We particularly thank David Matusik (FMNH) for obtaining the Putumayo specimen of P. werneri for AMNH, and A. M. Young (MPM) for providing a Choco specimen for dissection. For location of additional material we thank David Matusik, H. J. Hannemann (ZMH), and three anonymous private collectors. The following also assisted in efforts to locate specimens: Philip Ackery (BMNH); L. D. Miller (AME); R. dejong (RMNH); J. E. Rawlins (CMNH); and K. S. Brown Jr (various South American inquiries). F. H. Rindge (AMNH) kindly facilitated assistance at AMNH including obtaining archival material. Robert Aronheim (Oakton, Virginia) generously provided an AMNH patron grant in 1981 for initial study of Agrias and Prepona butterflies which was subsequently followed by a similar grant from Joel B. Grae (Harrison, New York).
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Received for publication 1 April 1987; accepted 27 May 1988.