Research Article |
Corresponding author: Shinichi Nakahara ( snakahara@fas.harvard.edu ) Academic editor: Carlos Peña
© 2015 Shinichi Nakahara, Mario Alejandro Marín, Cristóbal Ríos-Málaver.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Nakahara S, Marín MA, Ríos-Málaver C (2015) Taxonomic status and redescription of Magneuptychia nebulosa (Butler, 1867) (Lepidoptera, Nymphalidae, Satyrinae) with a lectotype designation. ZooKeys 503: 135-147. https://doi.org/10.3897/zookeys.503.9156
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A redescription of Magneuptychia nebulosa (Butler, 1867), a poorly known euptychiine butterfly, is given here, and accurate distributional data are provided for the first time. Taxonomic status of this taxon has been discussed by comparing its morphology against its possible congeners. In addition, lectotype designation for M. nebulosa is provided in order to objectively establish the identity of this taxon and consequently stabilize the nomenclature.
Se presenta una redescripción de Magneuptychia nebulosa (Butler, 1867), una mariposa euptychina poco conocida y se brinda por primera vez datos de su distribución. Se discute el estatus taxonómico de este taxón por comparación de su morfología con la de posibles congéneres. En adición, se provee un lectotipo para M. nebulosa, en búsqueda de establecer objetivamente la identidad de este taxa y consecuentemente estabilizar su nomenclatura.
Euptychiina, Neotropical, Taxonomy, Venezuela, Morphology
Euptychiina, Neotropical, Taxonomía, Venezuela, Morfología
Magneuptychia Forster, 1964 is one of the most diverse genera in the subtribe Euptychiina (Lepidoptera: Nymphalidae: Satyrinae: Satyrini), containing 32 described species and many undescribed species (
This paper focuses on the scarce and poorly known species Magneuptychia nebulosa (Butler, 1867). This specific epithet was previously found to have been misapplied to other taxa and almost no accurate information was available regarding the species’ taxonomy, biology, and distribution. Therefore, a redescription of M. nebulosa based on a morphological analysis is provided, enabling future researchers to confidently identify this taxon. Two related taxa, Magneuptychia mimas (Godman, 1905) and Magneuptychia alcinoe (C. & R. Felder, 1867) were also studied in details and directly compared to M. nebulosa. The first accurate locality data for M. nebulosa is also provided. During our examination of Magneuptychia specimens, we found that the M. nebulosa type label was misapplied. Consequently, a lectotype designation for M. nebulosa is included in order to objectively establish the identity of this taxon and stabilize the nomenclature.
Morphology. Male and female genitalia were dissected using the methods of
AN Andrew Neild collection, London, UK
BMNH The Natural History Museum, London, UK
IVIC The Venezuelan Institute for Scientific Research, Miranda, Venezuela
MGCL McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, Florida, USA
Photographs of additional specimens (
Magneuptychia nebulosa: LECTOTYPE: Male (Fig.
Magneuptychia mimas: SYNTYPE: Male (Fig.
Magneuptychia alcinoe: SYNTYPE: Female (Fig.
Magneuptychia adults (dorsal view on left, ventral view on right): a Male M. nebulosa from Miranda, Venezuela b Female M. nebulosa from Miranda, Venezuela c Lectotype of M. nebulosa (photo credit: Trustees of the Natural History Museum, London) d Syntype of M. alcinoe (photo credit: Trustees of the Natural History Museum, London) e Syntype of M. mimas (photo credit: Trustees of the Natural History Museum, London).
Magneuptychia alcinoe: Male, Ecuador: Tunguruhua Prov., Rio Machay, 1700m July 4–5 1993 J. P. W. Hall & K. R. Willmott (MGCL) KW-13-05; Male, Ecuador: Pichincha Prov., Rio Toachi, Union del Toachi 800m June 30 1993 J. P. W. Hall & K. R. Willmott (MGCL) KW-13-06; Colombia: Antioquia, Municipio de Amalfi, bosque Picardia N6°47'33", W75°06'36", 1050msnm October 12 2007 09:30:00 borde de bosque A. M. Velez (MEFLG); Male, Ecuador: Tandapi, Rio Pilaton, Pichincha 1500m August 10 1993 J. P. W. Hall & K. R. Willmott (MGCL) SN-14-57; Female, Ecuador: Esmeraldas Prov. Rd. Lita-Alto Tambo km.16. 850m June 6 1994 J. P. W. Hall & K. R. Willmott (MGCL) SN-14-60; Female, Colombia: Valle, Bitaco, 1700m Jan 7 1985 J. B. Sullivan (MGCL) SN-14-107.
Euptychia nebulosa
MALE: forewing length 19.6–21.5 mm (n=4).
Wing shape. Forewing with costa slightly convex, inner margin straight, outer margin almost straight, medium section slightly concave, anterior slightly convex, angular. Hind wing rounded, slightly angular, base of costa convex, inner margin convex beyond vein 3A, tornus rounded, outer margin slightly undulating, apex slightly angular.
Wing venation. Forewing recurrent vein absent; hindwing humeral vein present.
Dorsal surface. Forewing ground color brown, submarginal band dark brown, undulating, extending from apex towards tornus, delimiting the slightly darker area, marginal band dark brown, extending from apex towards tornus, fringe greyish brown.
Hindwing color brown, submarginal band dark brown, undulating, extending from apex towards tornus, convex in each cell; marginal band dark brown extending from apex towards tornus; postmarginal and tornal areas pale ocher, fringe greyish brown.
Ventral surface. Forewing ground color chestnut brown, paler along inner margin; discal band thin, straight, reddish brown, extending from radial vein to just beyond vein 2A; postdiscal band reddish brown, weakly undulating, slightly wider than discal band, extending from radial vein and traversing towards inner margin until vein 2A, curved basally in cell Cu2-2A, approximately 2/3 distance from wing base to apex; faint band between postdiscal and submarginal bands dark brown, broad, extending from radial vein to just beyond vein Cu2; submarginal band dark brown, undulating, extending from apex to tornus, becoming less undulating towards the tornus, parallel to postdiscal band; marginal band dark brown, darker than submarginal band, almost straight, extending from apex towards tornus; narrow band distal to marginal band, dark brown, traversing outer margin, delimiting remaining area and fringe; submarginal ocellus in cell M1-M2 black with two white pupils and orange ring; fringe brown.
Hindwing ground color same as forewing, overlaid with subtle whitish pearly cast along inner margin and basal area; discal band reddish brown, slightly distally curved, extending from costal margin to inner margin, approximately 1/3 distance from wing base to apex; undulating postdiscal band color and width same as discal band, weakly undulating, traversing from costal margin towards inner margin, slightly bent basally in discal cell, curved distally in cell Cu1-Cu2 and curved inwards towards the anal margin below vein 2A, approximately 2/3 distance from wing base to apex; submarginal band dark brown, extending from apex towards tornus, curved basally in each cell; dark brown marginal band traversing along marginal line from apex towards tornus; narrow band distal to marginal band, band dark brown, traversing along outer margin, delimiting remaining area and fringe; five submarginal ocelli present, cells Rs-M1 and M1-M2 each with black, orange-ringed ocellus with two white pupils, M1-M2 ocellus relatively large (compared to ocellus in cell Rs-M1), cells M2-M3 and M3-Cu1 each with orangish relatively small ocellus, sometimes indistinct, Cu1-Cu2 with black, orange-ringed ocellus with two white pupils, similar in size to M1-M2 ocellus; fringe greyish brown.
Head. (Fig.
Magneuptychia nebulosa: a head in lateral view b male genitalia (SN-14-35) in lateral view c male genitalia in dorsal view d female genitalia (SN-14-59) in ventral view e female genitalia (SN-14-59) in dorsal view f female genitalia (SN-15-44) in dorsal view including sternites g male foreleg (tarsus, tibia and femur) h female foretarsus. Abbreviations: ae, aedeagus; br, brachia; cb, corpus bursae; db, ductus bursae; la, lamella antivaginalis; pa, papillae analis; sa, saccus; ds, ductus seminalis; te, tegumen; tg, tergite; un, uncus.
Legs. Tarsal segments reduced, foretarsus and tibia equal in length, femur slightly longer (Fig.
Abdomen. Eighth tergite and sternite reduced.
Genitalia. (Fig.
FEMALE: Similar to male except as follows: Wings wider and rounder; forewing length 19.6–21.0 mm (n=8); dorsal ground color slightly paler; ventral ground color pale ocher, discal and postdiscal bands orange-brown; foretarsus divided into 5 segments (Fig.
(Fig.
The female specimen in the AN collection is from a humid lower montane forest isolated on a ridge line along the southern slope of Cordillera de la Costa. The vegetation here has trees with a canopy over 15 m high, such as Miconia sp. (Melastomataceae), Palicourea sp. (Rubiaceae), Clusia sp. (Clusiaceae), and Chusquea sp. (Poaceae) (pers. obs.). Several true cloud forest inhabitants (e.g. Evenus coronata (Hewitson, 1865) (Lycaenidae), Corades enyo enyo (Hewitson, 1849) (Nymphalidae) and Epiphile epicaste epicaste (Hewitson, 1857) (Nymphalidae) are also recorded here (pers. obs.). Thus, it is reasonable to expect that M. nebulosa can also be found in lower cloud forests on the slopes of the Cordillera de la Costa.
Phenotypically, M. nebulosa most closely resembles M. alcinoe and M. mimas. These species can be distinguished from M. nebulosa by size and wing pattern. Magneuptychia nebulosa is relatively small and possesses a wavy, somewhat irregular postdiscal band (slightly bent basally in discal cell, curved distally in cell Cu1-Cu2) of the ventral hind wing, whereas M. alcinoe and M. mimas are larger and have a straight hindwing postdiscal band. This straight ventral hindwing postdiscal band is also present in all other Magneuptychia. However, some of the members of Paryphthimoides (e.g. P. poltys (Prittwitz, 1865)) also exhibits this curved postdiscal band. Magneuptychia nebulosa possesses a rather reddish discal and post discal bands. The number of white pupils in the five ventral hindwing subapical ocelli varies within M. alcinoe and is thus occasionally diagnostic; some specimens of M. alcinoe have only one pupil in one of the ocellus (K. Willmott, pers. comm.), whereas M. nebulosa always have two pupils in four ocelli, and one pupil in the larger, fifth ocellus. In addition, M. nebulosa may be confused with a variation of M. modesta (Butler, 1867), which is a species that seems to be very variable and is perhaps a complex of several species. However, M. nebulosa differs from this taxon by the combination of the undulating ventral hindwing postdiscal band and double-pupilled ocelli in ventral hindwing cell M1-M2 (usually one in M. modesta).
Magneuptychia nebulosa was described from Venezuelan specimens. The type series was originally deposited in the Dyson collection and subsequently purchased by the BMNH in 1847 (G. Lamas, pers. comm.), where it is now deposited. The extant type series consists of one male (Fig.
Butler provides a precise forewing measurement of 1.55 inches (39.37 mm). This theoretically makes it possible to deduce which specimen was the subject of Butler’s description, though the male and female syntypes have nearly identical forewing lengths of 39 mm and 40 mm, respectively. Butler’s description also clearly refers to five ocelli, three of which are relatively small with two pupils. Male ocelli have two pupils, but female ocelli only have one. Therefore, we designate the male specimen as the lectotype of M. nebulosa. This is important because: (a) this male specimen most closely agrees with the original description, (b) it is in better condition than the female specimen, and (c) the male genitalia of euptychiine species are better known and are more commonly figured than female genitalia, and therefore have more scope to delimit species. The specific epithet nebulosa has been incorrectly applied to different taxa in
The male and female genitalia of M. nebulosa are extremely similar to those of M. alcinoe. Despite dissecting several specimens per species, we could not find any convincing species-level differences except for their overall size, which appears to correlate to the differences in overall body size. However, the male genitalia exhibit some variation of the costal region and cucullus of the valvae, as well as variation in patterns of the cornuti. Further examination of these structures could provide diagnostic characters for male specimens of these taxa.
In general, most euptychiine species are distinguishable from their congeners by characters of the male and female genitalia, so the genitalic similarity would ordinarily suggest that M. nebulosa and M. alcinoe are conspecific. However, the small adult size of M. nebulosa, its rather reddish bands, its curved ventral hindwing post discal band, and its rather small ocelli are all consistent and appear to be reliable characters to distinguish it from M. alcinoe. We were not able to find records or specimens of M. alcinoe from Cordillera de la Costa, nor M. nebulosa from an area inhabited by M. alcinoe, suggesting that the two species are allopatric. Cordillera de la Costa is isolated from the adjacent Sierra de Turimiquire and Cordillera de la Mérida by flat and scrubby lowlands, namely the depression of Yaracuy and the depression of Unare, respectively (see Fig.
Although the status of M. nebulosa is currently resolved, other uncertainties about Magneuptychia remain. For example, the type of M. mimas closely resembles that of M. alcinoe, leading some to suggest that these taxa are conspecific, with the former merely being a Bolivian population of the latter. Conversely, these similarities may instead suggest the need for an “alcinoe” species group to distinguish these very similar taxa from other Magneuptychia. The relationships between M. nebulosa and its congeners are still not fully understood; a revision of the genus is crucial to facilitate identification of euptychiine species in museum collections. Once we have a better understanding of Euptychiina and can reliably identify them, they can be used in broader, higher-impact studies of conservation and biogeography.
We thank Andrew Neild (UK) for discussion and for providing photos and valuable comments on the manuscript. Thanks are also due to Gerardo Lamas (Peru) for valuable discussion throughout the course of preparing this article; to David Lees (BMNH) and Blanca Huertas (BMNH) for kindly dissecting specimens at BMNH and for their help; to BMNH for allowing us to use photos, and to David Plotkin (MGCL) for correcting English and for providing comments on the manuscript. We are also grateful to Colciencias (grant No. 1118-521-28378) and the National Science Foundation (grant No. DEB-1256742).
Records for M. nebulosa from Quebrada Honda, El Jarillo, Miranda, Venezuela
Data type: Adobe PDF file
Explanation note: Records for M. nebulosa from Quebrada Honda, El Jarillo, Miranda, Venezuela; Altos de Pipe, Instituto venezolano de Investigaciones Científicas, Miranda, Venezuela: Cristóbal Ríos Málaver Leg: These following specimens are deposited in the reference collection of the Venezuelan Institute of Scientific Research IVIC, Altos de Pipe, Miranda, Venezuela.