ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia ISSN 1315-642X
Anartia, 28 (junio 2019): 26-46
M.G. Andrade-C., A.L. Viloria, E.R. Henao-Bañol & J.-F. Le Crom
http://zoobank.org/urn:lsid:zoobank.org:pub:E561593A-FC19-4512-B6D2-B6386360BF00
Reasignaciones genéricas de mariposas satírinas de Colombia y Venezuela (Lepidoptera: Nymphalidae)
M. Gonzalo Andrade-C.1, Ángel L. Viloria2, Efraín R. Henao-Bañol1,4
& Jean-François Le Crom3
1Instituto de Ciencias Naturales - ICN, Universidad Nacional de Colombia, Bogotá, Colombia.
2Centro de Ecología, Instituto Venezolano de Investigaciones Científicas (IVIC), km 11 carretera Panamericana, Altos de Pipe, Edo. Miranda, Venezuela.
3Calle 45 N° 9-42, ofic. 202, Edif. Carlec, Bogotá, Colombia.
4Instituto Amazónico de Investigaciones Científicas SINCHI. Calle 20, No. 5-44, Bogotá, D. C., Colombia
Correspondencia: M. G. Andrade-C.: mgandradec@unal.edu.co
(Recibido: 05-03-2018 / Aceptado: 08-11-2018)
ABSTRACT
Four genera of butterflies belonging to the Nymphalidae: Satyrinae are erected, diagnosized, described and discussed in or- der to reassign ten species recently classified within the genus Magneuptychia Forster, several of which occur in Colombia and Venezuela: Nubila Viloria, Andrade & Henao, gen. nov. (to include Euptychia gera Hewitson, E. metagera Butler, E. moderata Weymer, E. nortia Hewitson, E. purusana Aurivillius and E. segesta Weymer), Macrocissia Viloria, Le Crom & Andrade, gen. nov. (including Neonympha iris C. Felder & R. Felder and Euptychia inani Staudinger), Colombeia Viloria, Andrade & Le Crom, gen. nov. (including Euptychia mycalesis Röber) and Omacha Andrade, Viloria, Henao & Le Crom, gen. nov. (including Magneuptychia pax Huertas, Lamas, Fagua & Willmott). Additionally, Euptychia tiessa Hewitson is transfered to the genus Satyrotaygetis Forster, which is revised. Eleven new combinations are established, one new synonym is proposed and two species are reinstated.
RESUMEN
Se erigen, diagnostican, describen y discuten cuatro géneros de mariposas pertenecientes a las Nymphalidae, Satyrinae, con el fin de reubicar diez especies recientemente clasificadas dentro del género Magneuptychia Forster, varias de las cuales se encuentran en Colombia y Venezuela: Nubila Viloria, Andrade & Henao, gen. nov. (para incluir Euptychia gera Hewitson,
E. metagera Butler, E. moderata Weymer, E. nortia Hewitson, E. purusana Aurivillius y E. segesta Weymer), Macrocissia
Viloria, Le Crom & Andrade, gen. nov. (incluyendo Neonympha iris C. Felder & R. Felder y Euptychia inani Staudinger), Colombeia Viloria, Andrade & Le Crom, gen. nov. (incluyendo Euptychia mycalesis Röber) y Omacha Andrade, Viloria, Henao & Le Crom, gen. nov. (incluyendo Magneuptychia pax Huertas, Lamas, Fagua & Willmott). Adicionalmente se transfiere Euptychia tiessa Hewitson al género Satyrotaygetis Forster, el cual se revisa. Se establecen once nuevas combina- ciones, se propone un nuevo sinónimo y se restituyen dos especies.
Generic reassignments of satyrine butterflies
INTRODUCTION
The apparent polyphyletic and paraphyletic natures of the genus Magneuptychia, as loosely diagnosized by Forster (1964), has been pointed out by Murray & Prowell (2005: 72), Marín (2011: 13, 84); Marín et al. (2011: 9, 10; 2017:
768, 774, 781, 182) and Costa et al. (2016: 200). The lat- ter authors discussed this historical case in some detail and designated a Neotype for Papilio libye Linnaeus, the type species of Magneuptychia Forster. This nomenclatural act, followed by a comparative study of morphology and the consideration of genetic evidence available through phy- logenies recently published by molecular scientists, al- lowed for the establishment of the boundaries of the genus based on morphological singularities of its type species and the application of basic logic. Thus, Costa et al. (2016) recognized only three nominal species within the genus:
M. libye, M. libyoidea Butler and M. lethra Möschler (plus one yet to be described), and explicitly excluded 36 taxa from Magneuptychia. Notwithstanding the foregoing, re- cent work by a number of experts has deliberately chosen to disdain such taxonomic proposal (Huertas et al. 2016, Marín et al. 2017, Zacca et al. 2017), even if their results may appear contradictory with their approaches.
In inventorying and classifying the satyrine butterfly fauna of our countries, Colombia and Venezuela, we have dealt with the generic allocation of a number of species for- merly assigned to Magneuptychia. In this article we erect, di- agnosize, describe and discuss four genera belonging to the Nymphalidae, Satyrinae, in which we reassign ten species: Nubila Viloria, Andrade & Henao, gen. nov. (for Euptychia gera Hewitson, E. metagera Butler, E. moderata Weymer,
E. nortia Hewitson, E. purusana Aurivillius and E. segesta Weymer), Macrocissia Viloria, Le Crom & Andrade, gen. nov. (for Neonympha iris C. Felder & R. Felder and Eupty- chia inani Staudinger), Colombeia Viloria, Andrade & Le Crom, gen. nov. (for Euptychia mycalesis Röber) and Oma- cha Andrade, Viloria, Henao & Le Crom, gen. nov. (for Magneuptychia pax Huertas, Lamas, Fagua & Willmott). Additionally, we transfer Euptychia tiessa Hewitson to the genus Satyrotaygetis Forster, which is revised.
MATERIALS AND METHODS
Apart from collateral biological evidence gathered from published sources and considered in the discussion of this article to support our systematic hypotheses, this study is solely based upon the common practices of comparative morphology: observation, recognition and interpretation of homologies. The resulting taxonomy constructed with the information thus generated obeys logical inference of
set theory. We classify in the same category one or more species that share similar and equivalent morphological traits. Each group of species embraced by the four entities herein treated as new genera share common distinctive phenotypic characters in wing venation and wing visual pattern, as well as in the shape and proportions of certain major structures of the male genitalia. The sharing of those morphological traits appears sufficiently informative for our purposes and makes cohesive each of the proposed species groups. Therefore, their taxonomy coherent and plausible, but by no means exempt of further test by ap- propriate modern genetic techniques.
In this work we recognize fourteen species and allocate them in five genera. In the past, those species had been described as, or arbitrarily ascribed, to Euptychia Hübner (1818), Splendeuptychia Forster (1964) and/or Magneup-
tychia Forster (1964).
Members of Euptychia, as defined in modern taxonomy exhibit some, possibly unique, morphological characters (Singer et al. 1983, Murray 2001, Viloria 2007, Neild et al. 2014), that are entirely absent in our four new genera.
In part for this reason, morphological diagnoses for the new genera established in this paper are almost entirely based on comparisons with Splendeuptychia and Mag- neuptychia. In few cases we exceptionally invoke characters distinctive for the genera Cissia Doubleday (1848), Saty- rotaygetis Forster (1964), Huberonympha Viloria & Costa (in Costa et al. 2016) and Stevenaria Viloria et al. (in Costa et al. 2016). However, elementary structures of the wing pattern characteristic for all ten genera are abstracted in comparative Table 1. In our opinion other satyrine but- terfly genera known in the American continent bear not enough resemblance to the taxa herein treated and they have not been taken into consideration.
Nomenclatural acts in this work follow the provisions of the latest edition of the International Code of Zoologi- cal Nomenclature (ICZN 1999).
Technical procedures
Morphological examinations were performed on dry museum specimens kept in standard entomological cabi- nets. As indicated for each case, they are in the custody of individuals or institutions mentioned in the abbreviations section, below. When necessary for study, a limited number of samples were dissected, chemically treated according to the purpose by following well known protocols, and their structures cleaned under magnification with appropriate microdissecting tools by standard methods and preserved either in liquid (ethanol/glycerin for genitalia) or in fixing media (euparal in glass slides for diaphanized wings). Most of this entomological material had been collected by some
M.G. Andrade-C., A.L. Viloria, E.R. Henao-Bañol & J.-F. Le Crom
of the authors in multiple expeditions to several natural re- gions of Colombia and Venezuela. In those cases the but- terflies were set, labeled, dissected and generally studied and photographed locally by each author. When necessary we have used magnifying optic devices and various sources of artificial illumination, like several brands and models of stereomicroscopes (Leica, Wild, Olympus) with coupled cameras, both lucid and photographic, for producing pre- cision drawings and colour pictures, respectively. Manual 35 mm digital photographic cameras have also been used to record all specimens examined and their data labels.
Abbreviations
comb. nov.: combinatio nova; e. g.: exempli gratia; FWL: Forewing length; gen. nov.: genus novo; gen. rev.: genus revised; ICN: Instituto de Ciencias Naturales, Uni- versidad Nacional de Colombia, Bogotá D.C., Colombia;
i. e.: id est; IVIC: Instituto Venezolano de Investigaciones Científicas, Centro de Ecología, Altos de Pipe, Venezu- ela; JFLC: Collection of Jean-François Le Crom, Bogotá, Colombia; MALUZ: Museo de Artrópodos, Facultad de Agronomía, La Universidad del Zulia, Maracaibo, Venezu- ela; MC: Collection of Mauro Costa, Caracas, Venezuela; MHNUC: Museo de Historia Natural, Universidad de Caldas, Manizales, Colombia; MIZA: Museo del Instituto de Zoología Agrícola, Facultad de Agronomía, Universidad Central de Venezuela, Maracay, Venezuela; NHMUK: The Natural History Museum, London, United Kingdom, MNKB: Museum für Naturkunde, Berlin, Germany, SMTD: Staatssammlung Museum für Tierkunde, Dres- den, Sajonia, Germany; ZSM: Zoologisches Staatssam- mlung München, Munich, Germany; sp. restit.: speciebus restituit; syn. nov.: synonymum novo; TL: Type locality.
RESULTS
Nubila Viloria, Andrade & Henao, gen. nov.
(Figs. 1, 2, 10, 11, 12)
http://zoobank.org/urn:lsid:zoobank.org:act:5C8ACA7D-8F2D-4FEB- BC57-984013C375C5
Type species: Euptychia gera Hewitson, 1850; herein
designated.
Diagnosis
Butterflies of small to medium size (FWL: 17.5-23 mm), superficially comparable in size and general aspect to
those of Splendeuptychia Forster sensu stricto*. However, it differs from the latter by having two well defined forewing underside central bands in postbasal and discal positions (faint and thin, postbasal and postdiscal in Splendeupty- chia, the latter much more divergent towards the tornus). Hindwing ocellus prominent on Cu1, with two pupillae in Nubila, only one in Splendeuptychia. Differs from Mag- neuptychia Forster sensu stricto** by having a smaller size and by lacking the sixth ocellus on the anal margin on the hindwing underside. Forewing vein R2 emerges very close to the root of R3+R4+R5 at the anterior distal extremity of the discal cell (similarly in Cissia Doubleday, Colom- beia, gen. nov. and Macrocissia, gen. nov.); in Magneup- tychia, Satyrotaygetis Forster, Stevenaria Viloria et al. and Huberonympha Viloria & Costa, R2 emerges well before the end of the discal cell, closer to the root of R1; Sc ends at two fifths of the length of the Costa, as in Stevenaria (but approximately at the middle point of the Costa in Cissia, Magneuptychia, Satyrotaygetis, Huberonympha, Co- lombeia, gen. nov. and Macrocissia, gen. nov.). Forewing crossvein m1-m2, forming a distinct angle that points basad, from this angle emerges a short, but distinct recur- rent veinlet, into the discal cell. Both angle and veinlet missing in Cissia, Magneuptychia and Huberonympha, but present in Satyrotaygetis, Stevenaria, Colombeia, gen. nov. and Macrocissia, gen. nov. (see below). Male genitalia with saccus slender, about same length as uncus (similarly to Satyrotaygetis, which exhibits more robust saccus and uncus; slightly shorter than uncus in Macrocissia, gen. nov., notably shorter than uncus in Cissia, Magneuptychia, Splendeuptychia, Huberonympha and Omacha, gen. nov.; much longer than uncus in Stevenaria); valvae simply sub- triangular (similar to Splendeuptychia and Huberonympha, but not quite as elongated as in Magneuptychia; not sub- rectangular as in Cissia, not hooked as in Stevenaria, not spine-tipped as in Satyrotaygetis, Colombeia, gen. nov. and Macrocissia, gen. nov., not tricuspid at the tip as in Oma- cha, gen. nov.); aedeagus shorter than uncus, a character not shared with any of the nine genera herein compared with.
Description
Antennae relatively short, reaching to one third of the costa. Forewings subtriangular, apex and tornus rounded, hindwings suboval, outer margin slightly sinuous. Vena- tion (Figs. 1, 2): forewing subcosta notably swollen at
* It is our criterion that only its type species, Euptychia ashna Hewitson (1869), and Euptychia ambra Weymer (1911), strictly qualify to belong to Splendeuptychia Forster. Other species allocated in that genus by several authors, as arranged in Warren et al. (2012b) are in need of generic re- assignment.
** As re-defined by Costa et al. (2016).
Generic reassignments of satyrine butterflies
basal half, gradually diminishing until third quarter its length, ending at two fifths of costa; R1 independently emerging at distal quarter of discal cell, ending within third fifth of the costa; R2 independent, emerging much closer to root of R3-R5 than to R1; root of R3, R4 and R5 emerging from anterior extremity of discal cell; R5 end- ing at apex; R3 and R4 ending more or less equidistant from the end of R2 and R5; M1, M2, M3, Cu1 and Cu2 all independent, emerging from the discal cell; crossvein m1-m2 forming a distinct angle pointing basad; a short but distinct recurrent veinlet emerging from its angle in- side the discal cell; m2-m3 longer than m1-m2, slightly curved distad; root of Cubitals notably swollen at basal half, closing the cell at base with the swollen basal por- tion of Sc; A2 not inflated, independent, slightly sinu- ous at basal third; discal cell about half of wing length. Hindwing humeral present, but short and diffuse; all principal veins independent; Sc + R1 emerging at base of cell, ending at two thirds of costa; Rs ending at apex; A2 and A3 independent but their basal portions running to- gether for about one fifth of A3, then diverging ; m3-cu1 nearly half length of m1-m2, m2-m3 and cu1-cu2 about one and a half the length of m1-m2; discal cell about three fifths of wing length. Ocellar pattern as described in Table 1. Male genitalia (Figs. 10-12): tegumen dome- like, relatively small (shorter than uncus); base of uncus lanceolate, clearly distinguished from tegument; subunci well formed, almost as long as uncus, robust at base; vin- culum stylized but strong ; appendices angulares regular- ly developed but not very conspicuous; saccus flattened dorsoventrally, laterally slender, tubular, slightly curved upwards, shorter than uncus; valvae subtriangular, exter- nally covered by short setae, only finely irregular along the dorsal edge between the ampulla and the cucullus; aedeagus shorter than uncus, thickset.
Derivatio nominis
As an adjective Nubila means “clear” in Latin, but it also means “clouds” as a substantive. Both meanings evoke visual reminiscences of the white colour pattern of the males of Euptychia gera Hewitson, its sexually dimorphic, type species. Among its known congeners, appropriately defined here by other relations of homology, males of E. metagera Butler are the only representatives in the group to share a comparable whitish aspect.
Species belonging to Nubila
Nubila gera (Hewitson, 1850), comb. nov.
[Euptychia gera Hewitson, MS, pl. 63, fig. 4 (female)],
nomen nudum
Euptychia gera Hewitson, 1850: 439-440 [female] [TL: Amazon]
Neonympha gera (Hewitson); Herrich-Schäffer, 1865: 70.
Euptychia gera Hewitson; Westwood, 1850: pl. 63, fig.
4, 1851: 373; Butler, 1867a: 494; 1868: 35; 1877:
121; Kirby, 1871: 54; 1879: 134; Weymer, 1911:
215; Gaede 1931: 447; Lamas, 1969: 346; Lewis,
1973, 1987: 233, pl.58, fig. 2 (male); d’Abrera, 1988:
770, 771, row [8] figs. [1] (male dorsal), [2] (male [sic] dorsal)(misidentification of E. metagera Butler, female), row [9] figs. [1] (female dorsal), [2] (male
ventral), 874; 2001: 183, pl. 137, fig. 17 (male);
Overal & da Silva, 2002: 532; Shou et al., 2006: 84.
Euptychia hiemalis Butler, 1867a: 494 [male]. [TL: Amazons]
Euptychia hiemalis Butler; Butler, 1867b: 109, pl. 12, fig. 4; 1868: 35; 1877: 121; Kirby, 1871: 54; Druce,
1876: 213; Weymer, 1911: 215; Aurivillius, 1929:
158; Gaede, 1931: 451; d’Abrera, 1988: 789, 874.
Euptychia metaleuca distingüenda Bryk, 1953: 58-59 [female] [TL: Manaos, sv. Amaz.] (synonymy esta- blished by Lamas, 2004: 220)
[Magneuptychia gera (Hewitson); Forster, 1964: 126,
127 fig. 153 (male genitalia) (misidentification)]
Splendeuptychia distinguenda (Bryk); Forster, 1964: 130.
Magneuptychia gera (Hewitson); Brown, 1991: 376; T.
& L: Racheli, 2001: 329; Brévignon, 2008: 73, 82, 84, 93, pl. 4, figs. 39a, 39b (male genitalia), 39c; Pas- cal et al., 2015: 108. 111 fig. F (male); Huertas et al., 2016: 7.
Magneuptychia gera gera (Hewitson); Lamas, 2004: 220; Rodríguez & Miller, 2013: 210.
M.[agneuptychia] gera (Hewitson); Costa et al., 2016: 199.
[¿?] gera Hewitson; Costa et al., 2016: 204. [¿?] hiemalis Butler; Costa et al., 2016: 204.
Nubila metagera (Butler, 1867), comb. nov.
Euptychia metagera Butler, 1867a: 494 [male]. [TL: Ega, U. Amazons]
Euptychia metagera Butler; Butler, 1867b: 109, pl. 11, fig. 4; 1868: 35; 1877: 121; Kirby, 1871: 54; Wey-
mer, 1911: 215; Gaede, 1931: 454; d’Abrera, 1988:
770, 771, figs. [1] (male syntype, dorsal), [2] (male
ventral), 875.
Euptychia nortia abrahami Bryk, 1953: 63 [female] [TL: Tarocuá, Sv. Amaz.] (synonymy established by Lamas, 2004: 220); Costa et al., 2016: 199 (as a sy- nonym)
M.G. Andrade-C., A.L. Viloria, E.R. Henao-Bañol & J.-F. Le Crom
Splendeuptychia abrahami (Bryk); Forster, 1964: 130.
[Euptychia gera Butler; d’Abrera, 1988: 771, row [9], fig. 2 (female dorsal misidentification)]
Magneuptychia metagera (Butler); Lamas, 2004: 220;
Rodríguez & Miller, 2013: 210.
E.[uptychia] metagera (Butler); Costa et al., 2016: 199. [¿?] metagera Butler; Costa et al., 2016: 204.
Nubila moderata (Weymer, 1911), comb. nov.
Euptychia nortia Hewitson f. moderata Weymer, 1911: 214, pl. 48 row g. [female] [TL: Mapiri, Bolivia] Euptychia moderata Weymer; Aurivillius, 1929: 158;
Lamas, 1969: 346.
Euptychia nortia Hewitson ab. moderata Weymer; Gae- de, 1931: 457.
Splendeuptychia moderata (Weymer); Forster, 1964: 130 [type examined].
[Euptychia nortia Hewitson; d’Abrera, 1988: 771, row [1], fig. [5] (female dorsal) (misidentification)]
M.[agneuptychia] moderata moderata (Weymer); La- mas et al., 1991: 10.
M.[agneuptychia] moderata (Weymer); Robbins et al., 1996: 231.
Magneuptychia moderata (Weymer); Lamas, 2003: 203; 2004: 220; Murray & Prowell, 2005: 69,72, 73,
75, 76; Peña et al., 2010: 247, 250 fig. 2, 251 fig. 3,
253 fig. 4.
E.[uptychia] moderata Weymer; Costa et al., 2016: 199. [¿?] moderata Weymer; Costa et al., 2016: 204.
Nubila nortia (Hewitson, 1862), comb. nov.
Euptychia nortia Hewitson, 1862: pl. [44], n. 2, fig. 2 [male] [TL: Amazon]
Neonympha nortia (Hewitson); Herrich-Schäffer, 1865: 70.
Euptychia nortia Hewitson; Butler, 1867a: 494; 1868: 35; 1877: 121; Kirby, 1871: 54; 1879: 134; Stau-
dinger, 1886: tfl. 80; 1887: 225; Weeks, 1905: 29;
Weymer, 1911: 214, pl. 49, row a; Gaede, 1931: 457;
d’Abrera, 1988: 770, 771, row [1], figs. [3] (male dorsal), [4] (female dorsal, erroneously identified as male), [5](female dorsal, misidentification of E. mo- derata Weymer), row [2], figs. [4] (male ventral), [5] (female ventral, misidentified as male), 876; Shou et al., 2006: 85.
Euptychia nortia Hewitson f. nobilis Weymer, 1911: 214, pl. 49 row a, syn. nov. [female] [TL: Tabatinga, Amazon, Brazil]
E.[uptychia] nortia Hewitson; Aurivillius, 1929: 158.
Euptychia nortia Hewitson ab. nobilis Weymer; Gaede, 1931: 457.
Magneuptychia nortia (Hewitson); Forster, 1964: 126,
127 fig. 152 (male genitalia), 130; Lewis, 1973,
1987: 233, 234, pl. 60, fig. 3 (female, nec male); Cos-
ta et al., 2016: 199.
Splendeuptychia nobilis (Weymer); Forster, 1964: 130.
Magneuptychia gera nobilis (Weymer); Lamas, 2004: 220.
Magneuptychia gera nortia (Weymer); Lamas, 2004: 220.
E.[uptychia] nobilis Weymer; Costa et al., 2016: 199. [¿?] nobilis Weymer; Costa et al., 2016: 204.
[¿?] nortia Hewitson; Costa et al., 2016: 204.
Nubila purusana (Aurivillius, 1929), comb. nov.
Euptychia purusana Aurivillius, 1929: 158, fig. 1 [male]. [TL: Rio Purus, Amazon]
Euptychia purusama [sic] Aurivillius, 1929: 158.
Euptychia purusana Aurivillius; Gaede, 1931: 462. [Euptychia ? segesta Weymer; d’Abrera, 1988: 771, row
[3], fig. [2] (male ventral, misidentification)]
S.[plendeuptychia] purusana (Aurivillius); Lamas et al., 1991: 10; Robbins et al., 1996: 232.
Splendeuptychia purusana (Aurivillius); Lamas, 2004: 222; Brévignon, 2008: 80, 82, 84, 94, pl. 6, figs. 65a, 65b (male genitalia), 65c; Peña et al., 2010: 247, 250 fig. 2, 251 fig. 3, 253 fig. 4.; Brévignon & Benmes-
bah, 2012: 52 (Brévignon 2008 and the last proba- bly misidentifications of an undescribed taxon).
Nubila segesta (Weymer, 1911), comb. nov.
Euptychia segesta Weymer, 1911: 214-215, pl. 51, row b [male] [TL: Gramal near Muzo, Colombia]
Euptychia segesta Weymer; Apolinar, 1928: 14; Gae- de, 1931: 464; d’Abrera, 2001: 183, pl. 137, fig. 18
(male); Shou et al., 2006: 85.
Euptychia ? segesta Weymer; d’Abrera, 1988: 770, 771, row [3], figs. [2] (male ventral) (misidentification of
E. purusana Aurivillius), [3] (male ventral), 877.
M.[agneuptychia] segesta (Weymer); Robbins et al., 1996: 231.
Magneuptychia segesta (Weymer); Lamas, 2004: 220. E.[uptychia] segesta Weymer; Costa et al., 2016: 199. [¿?] segesta Weymer; Costa et al., 2016: 204.
Weymer’s treatment of the species herein assigned to Nubila, gen. nov., is partly misleading. He did not illus- trate any sex of either Euptychia gera (= E. hiemalis) nor
E. metagera (= E. nortia abrahami). In the first case, he correctly identified the female, whose pattern descrip- tion corresponds well with that of the female syntype of Euptychia gera at the NHMUK (examined), but wrongly
Generic reassignments of satyrine butterflies
described the male of Euptychia metagera (syntype exam- ined, NHMUK) under the name of E. gera Hewitson. On the other hand, his description of E. hiemalis appears cor- rect, but he did not notice that it is in fact the strongly di- morphic male of E. gera. The latter name has 17 years pre- cedence over the first. The description of E. metagera given by Weymer also matches well the taxon described origi- nally by Butler under that name. We believe that Weymer also made a mistake in describing a female for Euptychia nortia Hewitson that does not correspond to that spe- cies (p. 214). Its illustrations (pl. 49 row a) appear much closer to E. purusana Aurivillius, but might represent an undescribed taxon from French Guyana, whose male is identified and illustrated as Splendeuptychia purusana (Aurivillius) by Brévignon (2008: 80, 84, pl. 6, figs. 65a, b, c). Weymer may have referred to the female he illustrated when mentioning Cayenne as a locality for E. nortia. We do not know of any reliable record of nortia for the Guy- ana region. The female of Nubila nortia, comb. nov., cor- responds to what is described by Weymer as form nobilis, syn. nov. (1911: 214, pl. 49, row a). It is also appropriately illustrated by Lewis (1973, 1987: pl. 60, fig. 3) and in the photograph by K. Garwood of one specimen from the sur- roundings of Cacaulandia (Rancho Grande) in the state of Rondonia, Brazil (under Magneuptychia gera nortia in Warren et al. 2012).
The male genitalia illustrated by Forster (1964: 127, fig. 153; herein reproduced in our Fig. 11) for “Magneupty- chia” gera does not correspond to this taxon, although it appears to represent another species of Nubila, gen. nov. We herein illustrate both male genitalia (Fig. 10) and wing venation (Fig. 1) for this species, the type of the genus.
Material examined
Nubila gera: BRAZIL: 1 female, Amazon, Hew. Coll.
79-69, B. M. Type No. Rh. 3187, Euptychia gera Hew., Type [Syntype], 1 male, Amazons, coll. By Wallace, 51- 119, B. M. Type No. Rh. 3186, Euptychia hiemalis Butl., Type [Syntype] (NHMUK); COLOMBIA: Amazonas: 2 males, La Pedrera, Río Apaporis, 165 m, 9.xi. 1994, leg. G. Andrade-C., GAC 6678, 6680, ICN-MHN-L 12634,
12635; Caquetá: 1 female, Araracuara, Río Mesay, Puerto Abeja, 450 m, 27.i.1998, leg. M. Porras, MP 171, ICN- MHN-L 24465; 2 males, 1 female, 13.i.1998, leg. M. Por- ras, MP 26, 20, 30, ICN-MHN-L 223724, 11311, 24466,
Gen. 1978; 2 females, San Vicente del Caguán, Caño Ima, 4.viii.1992, leg. J. Langley, ICN-MHN-L 12040, 12039;
2 females, San Vicente del Caguán, Caño Ima, 4.viii.1992, leg. J. Langley, ICN-MHN-L 12039, 12040; 1 female, San Vicente del Caguán, Mamachi, 21.vi.1992, leg. L. Langley, ICN-MHN-L 9047; Guainía: 1 male, 1 female, Inirida, La
Ceiba, 3,6329720 N, -67,8894720 W, 103 m, 21.iii.1998, leg. A. Gabanzo, ICN-MHN-L 23723, 35990 (ICN);
-72.876111 W, 240 m, 17.ii.2017, leg. E. Henao, EH 95, 219, ICN-MHN-L 18928, 18929 (ICN); 1 male San José
de Guaviare, i.[19]92, leg. J. Urbina?; 1 male, same data,
xi. [19]91; 1 male, same data, ix.[19]92; 2 males [1 Ge- nit. JFL 603] ( JFLC); Vaupés: 1 male, Taraira, Estación Caparú, 2.ix.[19]93+, leg. J. Pinzón ( JFLC); 1 female, Cararu, La Libertad, 300 m., 5.ix. 1993, leg. G. Fagua, ICN-MHN-L 12340; 1 male, Pacoa, Quebrada Jotabeya, Comunidad Jotabeya -0,6147220 S, -70,1897777 W, 328 m., 27.iii.2009, leg. L. González, ICN-MHN-L 25082; 1 female, Pacoa, Rio Apaporis, comunidad de Jirijirimo,
-0,4400000 S., -70,9607770 W., 360 m., 21.iii.2009, leg.
L. González, ICN-MHN-L25079; 1 female, same data, 19.iii.2009 ICN-MHN-L 25080; 1 female, same data, 21.iii.2009, ICN-MHN-L 25081; 1 female, same data, 27.iii.2009, ICN-MHN-L 25108; 1 male, Pacoa, Que- brada Jotabeya, Comunidad Buenos Aires -00°01’21,3” S,
-71°00’09,4” W, 181 m., 21.ii.2018, leg. E. Henao; 1 male,
Soratama – Río Apaporis, 250 m, iii.1952, leg. L. Richter, ICN-MHN-L 18881; 1 male Serrania de Chiribiquete bajo el Tepui, 24.viii.1994, leg. J. Langley, ICN-MHN-L 18889; VENEZUELA: 1 male, Amazonas, Morganito, 21.vii.2012, col. Mattei [Wing prep. ALV014-17 (fig. 1)]; 1 female, same locality, 6.xi.2014 [Genit. prep. ALV-566- 17]; 1 female, same locality, 11.x.2015; 3 males [1 Genit. prep. ALV-521-14 (fig. 10)], Amazonas, Mcpio. Guainía, camino Yavita-Maroa, 2° 55’16’’ N, 67° 26’25’’ W, 300 m, 29-31.viii. 1996, cols. J. Camacho, A. Heenan (MALUZ); 2 males, 1 female, Amazonas, Raudal del Danto, 5°02’39’’ N; 67°33’40’’W, 100 m, 19.xii.2016, M. Costa (IVIC).
Nubila metagera: BRAZIL: 1 male, Ega, U. Amazons,
H. W. Bates, Godman-Salvin coll. 1904-1, Euptychia metagera Btl., B. M. Type No. Rh. 3188, Type [Syntype] (NHMUK); COLOMBIA: Amazonas: 1 female, La Pedrera, 1 Kilómetro arriba trocha centro Providencia, 205 m, 16.xi.1994, leg. G. Andrade-C. GAC 6770, IC-
N-MHN-L 12871; Caquetá: 2 males, San Vicente del Caguán, Morichal Caño Rico, forest path, leg. J Langley ICN-MHN-L 9045, 9046 [Wing prep. ALV052-17 (fig.
2), Genit. prep. ALV568-17]; Vaupés: 1 male, Caparu, camino a mina “La Libertad”, 290 m, 29.viii.1993, leg. G. Fagua, G10-0175, ICN-MHN-L 12329; 1 female, Pacoa, Río Apaporís, Comunidad de Jirijirimo, -0,4400000 S,
-70,960777 W, 350 m, 19.iii.2009, leg. González-M., L.
A. leg. ICN-MHN-L 23083 (ICN); 1 female, Serranía de Chiribiquete, camino hacia el Tepui, leg. J. Langley, 31.vii.1994, ICN-MHN-L 18888; 1 female, Serranía de Chiribiquete, camino hacia el Tepui, leg. J. Langley,
M.G. Andrade-C., A.L. Viloria, E.R. Henao-Bañol & J.-F. Le Crom
7.viii.1994, ICN-MHN-L 18880; 1 female, Serranía de Chiribiquete, camino hacia el Tepui, 7.viii.1994, ICN
-MHN-L 18873; 1 female, Taraira, Estación Caparú, 2 sem. 93+, leg., J. Pinzón [Genit. JFL 612] ( JFLC); VE- NEZUELA: Amazonas: 1 male, Morganito, 11.x.2015, Col. Mattei (IVIC). Nubila moderata: BOLIVIA: 1 male, Mapiri, Collection v. Rosen, Cotypus Nr. Euptychia moderata Weym., Zoologische Staatssammlung Mün- chen (ZSBS). Nubila nortia: BRAZIL: Amazon: 1 male, Hewitson Coll. 79-69, Euptychia nortia Hew. 2, B. M. Type No. 3185, Euptychia nortia Hew., Type [Syntype] (NHMUK); 1 female, Brasilia, sept., Taperjos Präparat Nr. SA 108 Zoolog. Staatssmmlung München; 1 male Brasilia, sept., Rio Madeird, Manicore Jr., Amazonas Coll. Fassl in Coll Arp.; 1 male, Brasilia, sept., Tapajos viii. 20. Leg. Jr. Bug, Präparat Nr. SA 142 Zoolog. Staatssmmlun München; 1 male, Brasilia, sept. Tapajos, Amazonas, Coll. Fassl in Coll. Arp.; 1 male, Amazonas, Coll. Fassl in Coll, Arp., Präparat Nr. SA 109 Zoolog. Staatsam- mlung München (fig. 12, reproduced from Forster 1964) (ZSM); Nubila purusana: no specimens examined. Nubi- la segesta: COLOMBIA: Caldas: 1 female, Manzanares, 19.xii.1992, leg. J. Salazar [Genit. JFL 618] ( JFLC); 1 female, La Victoria, 1946, leg. E. Schmidt-Mumm, MHN-UC 303 (ICN); Tolima: 1 male, Lérida, vereda el Bledo, Finca El Bledo, 4, 8946100 N, -74,9696600 W, 9.viii.2009, leg. L. Ospina, 293 m, ICN-MHN-L 32239.
Macrocissia Viloria, Le Crom & Andrade, gen. nov.
(Figs. 3, 4, 13, 14, 15)
http://zoobank.org/urn:lsid:zoobank.org:act:879ECA3B-C455-478E-AF80- 1AB2FF9B7B7B
Type species: Neonympha iris C. Felder & R. Felder,
1867; herein designated.
Diagnosis
Butterflies of medium size (FWL: 22-23 mm) with as- pect comparable to that of the members of Magneuptychia sensu stricto, Satyrotaygetis and Colombeia, gen. nov. Dif- fers from Magneuptychia in that the females may exhibit dorsal ocellar elements on the hindwing, which however can be seen also in Satyrotaygetis and Colombeia, gen. nov. Macrocissia, gen. nov., has a single, forewing underside monopupillated ocellus (M1), like in Satyrotaygetis; it is bipupillated in Magneuptychia and Colombeia, gen. nov. Hindwing underside ocellar pattern widely differs from that of Satyrotaygetis in having ocelli R5, M1, M2 and M3 with two pupillae (monopupillated in Satyrotaygetis); Cu1 may have only one pupilla. Differs from Magneupty- chia in lacking the sixth microocellus over the anal margin. Hindwing dorsum of male Macrocissia exhibits darker an-
droconial scales, notably in M. iris (C. Felder & R. Felder), comb. nov., inner margin less excavated that in Magneup- tychia, Huberonympha and Colombeia, gen. nov. Wing ve- nation fundamentally differs from that of Magneuptychia by the presence of a vestigial recurrent vein into the discal cell, emerging from m1-m2 (similar to that in Satyrotay- getis and Colombeia, gen. nov., but less pronounced than in Nubila, gen. nov.). Male genitalia of Macrocissia bears distinctive valvae, with a hook-like distal process and a finely serrate cucullus. Similar characters are observable in species of Satyrotaygetis, where the cucullar and distal pro- cesses are however, separated by a constriction or diastema. A few other Neotropical species of satyrine butteflies with a similar shape in their valvae have been classified in the genera Pareuptychia Forster and Splendeuptychia, but their wing shape, size and patterns are not comparable to the species of Macrocissia, gen. nov.
Description
Antennae reaching to two fifths the length of the costa, shaft thin, club not conspicuous and gradually formed. Wings: forewing subtriangular, apex notably rounded, outer margin slightly convex, tornus moderately rounded. Hindwing suboval, outer margin softly scalloped, anal margin slightly excavated near the tornus. Venation (Figs. 3, 4): forewing as in Magneuptychia, except for the pres- ence of a short, primordial recurrent vein in m1-m2, and for the emergence of R1 closer to the base, at the begin- ning of the distal quarter of discal cell; roots of R5 and M1 closer to each other in Macrocissia, gen. nov. Male geni- talia (Figs. 13-15): origin of uncus more or less continu- ous with tegumen (unlike Magneuptychia); tegumen same length as uncus; gnathos well developed, emerging from both basal sides of the tegumen, almost reaching the tip of the uncus; vinculum robust, especially towards the pedun- culus; saccus tubular in lateral view, about the same length of gnathos; valvae subtriangular and elongated, reaching the tip of uncus, cucullus finely serrate with a distal hook- like distal spine pointing upwards; aedeagus robust, later- ally thicker than saccus, slightly curved upwards, about three quarters of the length of tegumen plus uncus.
Derivatio nominis
The undersides of the species we herein place within this genus are in external aspect reminiscent of Papilio clarissa Cramer (a junior synonym of Papilio penelope Fa- bricius), the type species of the genus Cissia Doubleday. However, they are considerably larger in wingspan. There- fore we name this new genus, Macrocissia, which literally means “long Cissia”, it is a feminine term compound by and adjective and a noun, and its etymological roots are
Generic reassignments of satyrine butterflies
Figures. 5-9. Wing venation of five species representative of four genera (two new) of Neotropical satyrine butterflies (males). 5. Colombeia mycalesis (Röber), comb. nov., type species of the genus Colombeia, gen. nov.; 6. Omacha pax (Huertas, Lamas,
3 4 Fagua & Willmott), comb. nov., type species of the genus Oma- cha, gen. nov.; 7. Satyrotaygetis tiessa (Hewitson), comb. nov.; 8.
Magneuptychia libye (Linnaeus), type species of the genus Mag-
Figures. 1-4. Wing venation of four species representative of two new genera of Neotropical satyrine butterflies (males). 1. Nubila gera (Hewitson), comb. nov., type species of the genus Nubila, gen. nov.; 2. Nubila metagera (Butler), comb. nov.; 3. Macrocissia iris (C. Felder & R. Felder), comb. nov., type species of the genus Macrocissia, gen. nov.; 4. Macrocissia inani (Stau- dinger), comb. nov.
neuptychia Forster; 9. Satyrotaygetis satyrina (H. W. Bates), type species of the genus Satyrotaygetis Forster.
Figures. 10-15. Lateral (and in last case ventral as well) views of the male genitalia structures of five species representative of two new genera of Neotropical satyrine butterflies. In lateral view drawings, left valve and aedeagus have been detached from arma- ture for better visualization. 10. Nubila gera (Hewitson), comb. nov., type species of the genus Nubila, gen. nov.; 11. Nubila
sp. [probably new, after Forster, 1964; not Magneuptychia gera (Hew.) as originally indicated]; 12. Nubila nortia (Hewitson), comb. nov. [after Forster, 1964]; 13. Macrocissia iris (C. Felder
& R. Felder), comb. nov., type species of the genus Macrocissia,
gen. nov. [after Forster’s Magneuptychia ayaya (= M. iris)], 1964;
14. Macrocissia inani (Staudinger), comb. nov., type specimen as illustrated by Forster, 1964 under Magneuptychia juani [sic]; 15. Macrocissia inani (Staudinger), comb. nov., upper lateral view, lower ventral view; detached aedeagus pointing to the left.
M.G. Andrade-C., A.L. Viloria, E.R. Henao-Bañol & J.-F. Le Crom
entirely Greek (μάκρος: long; Κισσία: a region of the an- cient Persian Empire).
Species belonging to Macrocissia
Macrocissia inani (Staudinger, [1886]), comb. nov.
Euptychia inani Staudinger, [1886]: tfl. 80. [TL: Rio S. Juan,Columb.]
Euptychia juani [sic] Staudinger, 1887: 226; Weymer, 1911: 214, pl. 49, row a; Gaede, 1931: 451; d’Abrera,
1988: 764, 765, row [2], figs. [1] (male dorsal), [2]
(male ventral), 875; Shou et al., 2006: 84.
E.[uptychia] juani [sic]; Staudinger; Schaus, 1913: 342. Magneuptychia juani [sic] (Staudinger); Forster, 1964: 126, fig. 150 (male genitalia, type); T. & L. Rache-
li, 2001: 329; Palacios & Constantino, 2006: 266;
Costa et al., 2016: 199.
[Euptychia] inani Staudinger; T. & L. Racheli, 2001: 329.
Magneuptychia inani (Staudinger); Lamas, 2004: 220;
Marín, 2011: 48 fig. 45, 56, 60, 65, 69, 71.
[¿?] inani Staudinger; Costa et al., 2016: 204.
Macrocissia iris (C. Felder & R. Felder, 1867),
Neonympha iris C. Felder & R. Felder, 1867: 483-484. [TL: Bogota]
Euptychia ayaya Butler, 1867a: 492-493, pl. 40, fig. 11. [TL: Tapajos, Amazons]
Euptychia ayaya Butler; Butler, 1868: 34; 1877: 122;
Kirby, 1871: 54; Holdich & Hinks, 1918: 234;
Gaede, 1931: 440; d’Abrera, 1988: 785, row [5], fig.
[1] (male dorsal), [2] (male ventral), 873; Lamas,
[1997]: 53 (synonymic priority established); 2004: 220 (as a synonym); T. & L. Racheli, 2001: 329 (as a synonym); d’Abrera, 2001: 183, pl. 135, figs. 34 male
dorsal, 35 male ventral; Brévignon, 2008: 74 (as a
synonym); Shou et al., 2006: 83.
Euptychia iris (C. Felder & R. Felder); Butler, 1868: 34, 1877: 122; Kirby, 1871: 54; Weymer, 1911: 221
(synonymy with E. agaya [sic] Butler established); Gaede, 1931: 440 (as a synonym).
Euptychia agaya [sic] Butler; Weymer, 1911: 221, pl.
49, row g; Chai, 1990: 56.
Euptychia agaya [sic] Butler f. azulina Ribeiro, 1931:
34. [TL: Jamary, Rondonia, Brasil]
Euptychia agaya [sic] azulina Ribeiro; Ribeiro, 1931: 40; May, 1933: 125 (synonymy established).
Magneuptychia ayaya (Butler); Forster, 1964: 126, 127
fig. 151 (male genitalia); Lamas, 1981: 29; Costa et
al., 2016: 199 (last one as a synonym of Neonympha iris C. & R. Felder ).
Euptychia agava [sic] Butler f. azulina Ribeiro; Mielke
& Casagrande, 1987: 22, 23, fig. 8.
M.[agneuptychia] ayaya (Butler); Lamas et al., 1991: 10.
M.[agneuptychia] iris (C. Felder & R. Felder); Robbins
et al., 1996: 231.
Magneuptychia iris (C. Felder & R. Felder); Lamas, [1997]: 53; 2004: 220; T. & L. Racheli, 2001: 329;
Brévignon, 2008: 74, 82, 84, 93, pl. 4, figs. 43a, 43b (male genitalia), 43c.
Chloreuptychia ayaya (Butler); Piñas, 2004: 35, figs.
279, 280.
Chloreuptychia agaya [sic] (Butler); Lewis, 1973, 1987: 233, pl. 55, fig. 17 (male); Shou et al., 2006: 82.
M.[agneuptychia] iris (C. Felder & R. Felder); Costa et al., 2016: 199.
[¿?] iris C. Felder & R. Felder; Costa et al., 2016: 204.
Judging by the similitudes of the structure of the val- vae of the male genitalia, and the homologies of most ele- ments of the wing pattern, the genus Macrocissia, nov., is close to Satyrotaygetis Forster and to Colombeia, gen. nov.
Material examined
Macrocissia inani: COLOMBIA: 1 male, Rio S. Juan,
Columb. [sic] Jr., ex collect. Staudinger, Origin., Eigen- tum Mus. Berlin, Präparat No. 149 Zoolog. Staatssam- mlung München (fig. 14, reproduced from Forster, 1964) [Lectotype Miller 1989]; 1 female Rio S. Juan, Columb. Jr., Origin., genitalia vial M-9093, [Paralectotype Miller, 1989] (ZMHB); Chocó: 1 male, Guadalupe, 50 m, leg. J. Umaña; 2 males, Tado, 20.iii.[19]90, leg. J. Salazar [Genit. JFL 606, (fig. 15)]; 1 female, Bahia Solano, 1.i.[19]97, leg.
J. F. Le Crom ( JFLC); Risaralda: 1 female Santa Cecilia, 650 m., x.1952, leg., J. Vélez, MHN-UC 13 (MHNUC).
prep. ALV571-17], 3050; 1 female, same data, xi.1959, leg. L. Richter, ICN-MHN-L 2708; 1 male, Santo Do- mingo, Santo Domingo de los Colorados, 500 m, iv.1959, leg. L. Richter, ICN-MHN-L 5980; 1 male, Pichincha, Sant[o] Domingo de los Colorados, 650 m, 25.10.1956, leg. J. Foerster (ZSM). Macrocissia iris: COLOMBIA: [Cundinamarca]: 1 male, Bogotá, Lindig, Type, iris, Fel- der Colln., Rothschild Bequest B. M. 1939-1 [Syntype of
Generic reassignments of satyrine butterflies
Neonympha iris C. Felder & R. Felder, label with note by
N. D. Riley: “Type of N. iris Feld. = E. ayaya Butl. Comp.
w. type xii.1912 NDR.”]; 1 male, Tapajos, Amazons, H.
W. Bates, B. M. Type No. 3264, Euptychia ayaya [male] Butl., Godman-Salvin Coll. 1904-1, Euptychia ayaya Butl. Type, H. T. [Holotype of Euptychia ayaya Butler, label with note by N. D. Riley: “Agrees with type of N. iris Feld. Brand perhaps rather bigger in this specimen, NDR.”] (NHMUK); 1 male, Butler Rio Guayabero, Ma- carena Sur, 300 m., 15.1.1959, leg. D. Trapp, label with note Abdomen gekilobt Haerelberm; 1 male Amazo- nas, Coll Fassl in Coll, Arp., Präparat Nr. SA 136 Zoo- log. Staatssmmlung München (fig. 13, reproduced from Forster, 1964) (ZSM), Caquetá: 1 female, San Vicente del Caguán, Caño Grande, 500 m, 17.viii.1992, leg. J. Langley, ICN-MHN-L 12043; 1 male, San Vicente del Caguán, Las Vegas, Pato Bajo, 500 m, 17.ix.1995, leg., G. Andrade-C., GAC 8037, ICN-MHN-L 24468 [Wing
prep. ALV055-17 (fig. 3), Genit. prep. ALV572-17)]; 2 males, same data, 12.ix.1995, leg, G. Andrade-C., GAC, ICN-MHN-L 14292, 37143; Guaviare: 1 male, San José del Guaviare, i.[19]92, leg. J. Urbina; 1 male, same data, ix.[19]92 (Genit. JFL 607)( JFLC); Meta: 1 male, San Juan de Arama, Serranía de la Macarena, 650 m, leg. L. Richter, ICN-MHN-L 37178; Vaupés: 1 female, Pacoa, Rió Apaporís, comunidad de Jirijirimo, 0.400000 N,
-70.9607770 W; 360 m, 20.iii.2009, leg. L. González, ICN-MHN-L 25390; 1 female, Soratama, Río Apaporís, i.1952, no more data, leg. L. Richter, ICN-MHN-L 6143; 4 females, mer. Umbria. SMD, (STMD-D); ECUADOR: 1 male, Ecuador, leg. Velástegui ( JFLC). SURINAM [?]: 3 male, SMD, (STMD-D).
Colombeia Viloria, Andrade & Le Crom, gen. nov.
(Figs. 5, 16)
http://zoobank.org/urn:lsid:zoobank.org:act:D2E62DCC-F456-4792- BC22-86BDA0801F29
Type species: Euptychia mycalesis Röber, 1927; herein
designated.
Diagnosis
Butterflies of medium size (FWL: 28-30 mm), superfi- cially comparable in size and aspect to the species belong- ing to Magneuptychia, Megeuptychia Forster and Satyro- taygetis. However, it is distinguished from these genera by the moderate but evident excavation of the anal margin near the tornus; never as soft as in Megeuptychia, nota- bly larger, or as marked as in Erichthodes Forster, which is smaller, but like in Huberonympha, from which it differs externally by never having sinuous or irregular underside dark bands. Butterflies belonging to Colombeia, gen. nov.,
bear double white pupillae on each complete ocellus of its underside (one pupilla in Satyrotaygetis, a combination of one and two or just one in Megeuptychia), not unlike Magneuptychia, which however is diagnosized by the pres- ence of a seventh micro-ocellus, at discal level on the anal margin. This ocellus is entirely absent in Colombeia, gen. nov., which is also characterized by the long diameter of its hindwing underside ocelli on cells M1 and Cu1 (round, never oval as in Magneuptychia). The latter especially con- spicuous. Magneuptychia and Omacha, gen. nov., both lack a recurrent veinlet emerging from m1-m2 in the fore- wing discal cell, a character common to Satyrotaygetis, Co- lombeia, gen. nov., Nubila, gen. nov. and Macrocissia, gen. nov. The last two genera, as well as Omacha, gen. nov., have characteristic elongated silver ocellar elements on hindwing underside cells M2 and M3. These features are missing in Colombeia. Due to the shape of the valvae, and the tegument-uncus configuration Colombeia, gen. nov., shows affinities in male genitalia to Magneuptychia. They however differ in wing pattern homologies as mentioned above.
Description
Antennae simple, reaching two fifths of the costa, club formed gradually and not conspicuous. Forewings sub- triangular, outer margin slightly excavated, apex and tor- nus moderately round. Hindwings suboval, outer margin crenulate, anal margin moderately excavated near tornus. Venation (fig. 5): Forewing subcosta notably inflated at basal third, root of cubitals moderately swollen at basal half; both converging at base to close the discal cell; A2 independent but closely attached to root of cubitals at base, slightly inflated in its first fifth of its length. Sc end- ing nearly at half length of costa; R1 emerging at the be- ginning of last distal quarter of discal cell, ending at limit of discal area; R2 independent emerging near tip of discal cell just before the root of R3+R4+R5; R3 emerging equi- distantly between extremity of cell and bifurcation of R4 and R5; R5 ending at apex; M1 to Cu2 all independent and running more or less parallel to the outer margin of wing; m1-m2 slightly curved inwards to cell; short recur- rent veinlet emerging from curvature, closer to root of M2; m2-m3 slightly curved outwards the cell. Hindwing Hu present, short, thick and diffused toward wing base; Sc ending at middle costa; RS ending at apex; all veins independent; A2 and A3 running closely together for the basal quarter of the second; m1-m2 just slightly concave; m2-m3 just slightly convex; m3-cu1 about half-length of m1-m2 and a quarter length of m2-m3. Ocellar pattern as presented in Table 1. Male genitalia (Fig. 16): Tegumen globular, origin of uncus differentiated from tegumen as in
M.G. Andrade-C., A.L. Viloria, E.R. Henao-Bañol & J.-F. Le Crom
Magneuptychia, but subunci emerging from the lower base of uncus, not as broad at the base as in Magneuptychia. Saccus as long as uncus, more conical and longer than that of Magneuptychia. Valvae subtriangular and elongated, reaching the same length as uncus, laterally less broad than those of Magneuptychia, but presenting the same kind of tiny processes at their distal third; never with serrate or spiny distal processes (like those in Macrocissia, gen. nov. or Satyrotaygetis). Aedeagus robust but more stylized than in Magneuptychia, lacking spiny areas.
Derivatio nominis
Colombeia is a word composed in Greek by the poly- math, cosmopolitan military leader and revolutionary Francisco de Miranda (Caracas, 1750-Cádiz, 1816), pre- cursor of the independence of Venezuela and Spanish America. It was intended to mean “the papers related to Colombia”, a name he gave to his monumental personal archives (63 volumes of documents, preserved in Caracas, Venezuela. http://www.franciscodemiranda.org/colombeia/), but it probably has no precise translation. Paraphrasing the Aristotelian Politeia, the name Colombeia derives from the notion of the “Great Independent State of Colombia” (Colombia is a latinization itself for the Columbian Conti- nent, honouring Christophorus Columbus). It was a uto- pian geopolitical project conceived by Miranda as early as 1798, as a single republic from the Mississippi to Cabo de Hornos. Simón Bolívar borrowed the name to establish Colombia (1819), a more modest Republic that, by 1822, embraced the current territories of Venezuela, Colom- bia, Panamá and Ecuador. Its dissolution began in 1830. Colombeia mycalesis Röber, comb. nov., is a distinctive butterfly species, so far the only member of its genus. Its distribution is restricted to the Western portion (biogeo- graphical Chocó) of the current Republics of Colombia and Ecuador (and possibly the adjacent Pacific slopes of Panamá). We think Colombeia, feminine substantive, is an appropriate name for a genus endemic to that region, and a tribute to Miranda.
Species belonging to Colombeia
Colombeia mycalesis (Röber, 1927), comb. nov.
Euptychia mycalesis Röber, 1927: 281. [TL: West-Co- lumbien, 500 meter]
Euptychia mycalesis Röber; Gaede, 1931: 456; d’Abrera, 1988: 764, 765 row [1], figs. [1] (male syntype ven-
tral), [2] (female dorsal), 875; 2001: 183, pl. 137,
fig. 7 (male); Lamas, 2001: 32; Shou et al., 2006: 85.
Magneuptychia mycalesis (Röber); Lamas, 2001: 32;
2004: 220; T. & L. Racheli, 2001: 329; Piñas, 2004:
34, figs. 267, 268; Palacios & Constantino, 2006:
266; Checa, 2008: 34 figs. (male ventral, dorsal). E.[uptychia] mycalesis Röber; Costa et al., 2016: 199. [¿?] mycalesis Röber; Costa et al., 2016: 204.
Material examined
Colombeia mycalesis: COLOMBIA: 1 male, West-
Columbien, 500 meter, März-Juni, mycalesis Type Röb [male], Brit. Mus. 1928-508, Type [Syntype](NHMUK); Chocó: 1 female, Río San Juan, 100 m, febrero 2009, leg.
J. Urbina [Genit. JFL 617]; 1 male, Rio St. Juan, Columb. Jr., Eupt. spec. 2, [genitalia vial 9091 Lee D. Miller]; 1 fe- male, Rio St. Juan. Columb. Jr. [genitalia vial 9092 Lee
Miller]; Valle del Cauca: 1 male, Bajo Calima, 300 m [Genit. JFL1859 (fig. 16)]; 1 female, same data ( JFLC); ECUADOR: 1 male, Santo Domingo, Santo Domingo de los Colorados, octubre 1959, ICN-MHN-L 28527 [Wing prep. ALV054-17 (fig. 5), Genit. prep. ALV550-17].
Omacha Andrade, Viloria, Henao & Le Crom, gen. nov.
(Figs. 6, 17)
http://zoobank.org/urn:lsid:zoobank.org:act:AC65B91F-7D82-40C7-93E9- DEED5D435F47
Type species: Magneuptychia pax Huertas, Lamas,
Fagua & Willmott, 2016; herein designated.
Diagnosis
Because of the superficial appearance, ocelli and bands on the underside, Omacha Andrade, Viloria, Henao & Le Crom, gen. nov., is comparable with other Neotropical genera as Magneuptychia Forster, Cissia Doubleday, Hu- beronympha Viloria & Costa, Stevenaria Viloria, Costa, Neild & Nakahara and Nubila Viloria & Andrade, gen. nov. Its type species herein designated is Magneuptychia pax Huertas, Lamas, Fagua & Willmott (in Huertas et al. 2016). However, the new entity differs homologically from all the genera mentioned in several characters of the wing size, pattern, venation and male genitalia. The pres- ence of a series of five, well developed, monopupillated ocellar elements from R5 to Cu1 on the hindwing verso, a character found also in the members of Nubila, which however have different venation and male genitalic struc- ture, is diagnostic for Omacha (see above).
Comparisons for the following description refer to Magneuptychia and Cissia sensu stricto, as considered by Costa et al. (2016)
Description
Butterflies of small to medium size (average FWL, male: ca. 21mm; female: ca. 20.5 mm). Antennae reach- ing two thirds of the costa, club formed gradually. Eyes
Generic reassignments of satyrine butterflies
prominent and setose. Palpi setose, twice as long as head, cream white in general, with white setae on the outer sides and dark brown ones in the inner, first segment subrect- angular in lateral view, second one cylindric, almost twice as long as first, third segment small and pointed. Fore- wings subtriangular, corners rounded, margins smooth; hindwings oblong, external margin slightly undulated. No androconial scales detectable. Ocellar elements well developed only in the hindwing: characteristically repre- sented by a series of five contiguous elements, with single pupils, in cells R5 to Cu1; those in R5, M1 and Cu1 are black ocelli fully developed; cells M2 and M2 bear ocel- lar elements lacking dense black around the pupils, but pupillar elements large and silvery (never elongate as in Nubila). Well developed ocellus in Cu1 above. Venation (Fig. 6): forewing Subcosta, R1 and root of Cubitals swol- len at base; Subcosta ending circa half length of the costa; R1 emerging at distal quarter of discal cell, ending within the beginning of distal third of the costa; R2 ending at costal limit of apex (at the end of fourth fifth of the costa in Magneuptychia, Huberonympha, Cissia and Stevenar- ia); R3 ending at tip of apex (before apex in Magneupty- chia, Huberonympha, Cissia and Stevenaria); R4 and R5 ending at outer margin limit of apex (R4 before apex and R5 at apical tip in Magneuptychia, Huberonympha, Cissia and Stevenaria); M1 emerging independently from Rs, like in Huberonympha and Stevenaria, but both roots not as close as in Magneuptychia, and much less distant from each other than in Cissia; root of A2 not swollen, emerg- ing independently and separate from discal cell (unlike other genera considered here for comparisons); m1-m2 strongly curved basally, resembling the case of Stevenaria, but without a recurrent veinlet towards the interior of the cell; m2-m3 sinuous. Hindwing Humeral well developed, reaching the coastal margin at the distal end of the first quarter of the coastal length, emerging together with Sc
+ R1 and Rs; A3 emerging from first sixth of A2 (unlike Magneuptychia, Huberonympha, Cissia and Stevenaria, in which it rises independently from the wing base); rs-m1 shorter than in Magneuptychia, Huberonympha, Cissia and Stevenaria; m1-m2 long and strongly curved inwards the cell, contrasting with Magneuptychia, in which this vein is straight (and only slightly curved in Huberonym- pha, Cissia and Stevenaria). Male genitalia (Fig. 17): Male genital armature stylized; tegumen dome-like, relatively small; uncus straight, lanceolate, one and half times lon- ger than tegumen; subunci thick at base, emerging later- ally in divergent angle, below the base of uncus, curving upwards and ending above the latter, more or less at the same height of tegumen; aedeagus stylized, as long as tegumen + uncus; saccus long and tubular in lateral and
superior view; valvae elongate, characterized by a diag- nostic, unique tricuspid tip.
Derivatio nominis
Omacha is the aboriginal name for the largest of all the species of river dolphins of South America, Inia geoffren- sis de Blainville, but it is also the name of a prize winning non-government environmental organization (Fundación Omacha) established in Colombia in 1993 for the pur- poses of developing research and conservation projects in the Amazonia. We dedicate the name of this new genus of Amazonian butterflies to this successful institution and to its President, our friend and colleague Dr. Fernando Tru- jillo for his devoted labour to conservation biology in Co- lombia and other countries of South America.
Species belonging to Omacha
Omacha pax (Huertas, Lamas, Fagua & Willmott, 2016), comb. nov.
[‘Euptychia’? sp.: d’Abrera, 1988: 786-787, row [7], figs.
[1] (male dorsal), [2] (female ventral)]
[Magneuptychia sp. n. 1; Lamas et al., [1997]: 65]
[Magneuptychia [n. sp.]; Lamas, 2004: 220 (no. 1416)]
[Magneuptychia sp. 3; Piñas, 2004: 34, figs. 271-272]
Magneuptychia pax Huertas, Lamas, Fagua & Will- mott, 2016: 6-10, figs. 1A (male), 1B (female), 2A-D (male abdomen & genitalia), 2E-G (female abdomen & genitalia), H-I (larval head capsule), 3 (distribution & habitat). [TL: [Colombia, Putuma- yo], n[ea]r. Villa Garzón, Los Naranjos [1°34’13’’N, 75°43’W]]
Material examined
Omacha pax: COLOMBIA: Amazonas: 1 male,
Puerto Nariño, ix.[19]94+, leg. F. Montero ( JFLC); 1 fe- male, Leticia, Caño Yaguarcaca, 150 m, 3.xii.1982, leg. J. Salazar, MHN-UC (MHNUC); 1 male, Puerto Nariño, Camino San Martín de Amacayacu, 105 m, -3,7761111 S, -70,3444444 W, 1.iv.2015, leg. G. Andrade-C., GAC 11988 ICN-MHN-L 34065 (ICN); Caquetá: 1 male,
San Vicente del Caguán, Río Yarí, Puerto Mosco, 217 m, 1,0849166 N., -74,2861111 W, 13.iii.2017, leg. E.
Henao, EH 293, ICN-MHN-L 37152; 1 male, same data, 1,1218611 N, -74,2918889 W, 15.iii.2017, leg.
Henao, EH 319, ICN-MHN-L 37151; 1 male, same data, 185 m, 0,1218333 N,-74,0932777 W, 20.iii.2017, leg. E. Henao, EH 521, ICN-MHN-L 31449 [Wing
prep. (Fig. 6), Gen. No. 1649 (Fig. 17)]; 1 female, same data, 420 m, 27.vii.1995, leg. G. Fagua, ICN-MHN-L 31449 (ICN).
M.G. Andrade-C., A.L. Viloria, E.R. Henao-Bañol & J.-F. Le Crom
Figs. 16-21. Lateral (and in some cases dorsal and/or ventral) views of the male genitalia structures of six species representative of four genera (two new) of Neotropical satyrine butterflies. In lateral view drawings, left valve and aedeagus have been detached from arma- ture for better visualization. 16. Colombeia mycalesis (Röber), comb. nov., type species of the genus Colombeia, gen. nov.; 17. Omacha pax (Huertas, Lamas, Fagua & Willmott), comb. nov., type species of the genus Omacha, gen. nov.; 18. Magneuptychia libye (Lin- naeus), type species of the genus Magneuptychia Forster (after Costa et al. 2016); 19. Satyrotaygetis incerta (Butler & Druce) (example from Costa Rica, after Hayward, [1964]. Not Euptychia satyrina (H. W. Bates) as originally indicated); 20. Satyrotaygetis gigas (Butler) (after Forster, 1964); 21. Satyrotaygetis tiessa (Hewitson), comb. nov.; upper lateral view, lower ventral view; aedeagus missing.
Generic reassignments of satyrine butterflies
Taxonomic position of Euptychia tiessa Hewitson, 1869
(Figs. 7, 21)
Molecular studies by Murray & Prowell (2005: 77) and Marín et al. (2017: 15) detected and drew the atten- tion to the apparent genetic closeness of Magneuptychia tiessa (Hewitson) and Satyrotaygetis satyrina (Bates) as shown by their resulting phylogenetic trees (sister species). Wing pattern of the members of Satyrotaygetis Forster is characterized by monopupillated ocelli unlike any other taxon treated in this work (see Table 1). This singularity is combined with a distinctly serrate cucullus, a hooked distal process of the valva, and a thick aedeagus (see corre- sponding correct figures of the male genitalia of S. incerta in Hayward, [1964]: 513, fig. 101, and S. gigas in Forster, 1964: 75, fig. 35; herein reproduced as Figs. 19 and 20, respectively). E. tiessa not only shares these external (habi- tus in Warren et al. 2012a) and internal (male genitalia in fig. 21, this work) characters, but also shows the typical venation (fig. 7) of other members of Satyrotaygetis (e. g., S. satyrina, type of the genus, as illustrated by Forster, 1964: 75, fig. 34; herein reproduced in Fig. 9). Genitalia and venation are both very different from those of the type species of Magneuptychia, Papilio libye Linnaeus (see figs. 8 venation, 18 male genitalia). This cumulative evidence allows for confidently transfer E. tiessa to Satyrotaygetis.
Satyrotaygetis Forster, 1964, gen. rev.
Satyrotaygetis Forster, 1964: 73; Miller, 1968: 94; La-
mas, 2004: 222; Costa et al., 2016: 200.
Type species: Taygetis satyrina H. W. Bates, 1865; by original designation (Forster, 1964: 73).
Species belonging to Satyrotaygetis
Satyrotaygetis gigas (Butler, 1867a), sp. restit.
Euptychia gigas Butler, 1867a: 486, pl. 40, fig. 7 [TL: México]
Euptychia gigas Butler; Butler, 1868: 30; 1877: 120;
Kirby, 1871: 52; Hayward, [1964]: 514; Lamas, 2004: 222 (last two as as synonym of S. satyrina (H.
W. Bates))
Euptychia satyrina (H. W. Bates) f. gigas Butler; Wey- mer, 1911: 212, pl. 48, row e.
Euptychia satyrina (H. W. Bates) var. gigas Butler; Gae- de, 1931: 464.
Satyrotaygetis gigas (Butler); Forster, 1964: 73, 75 fig.
35 (male genitalia), t. 29 fig. 7; Raguso & Llorente, 1997: 287.
Cissia gigas (Butler); DeVries, 1987: 275, pl. 48, fig. 6
(male), 7 (female); Shou et al., 2006: 82.
[Euptychia satyrina H. W. Bates; d’Abrera, 1988: 781, row [5] (male ventral, pale form) (misidentifica- tion)]
[Satyrotaygetis satyrina (H. W. Bates); Chacón & Montero, 2007: pls. 177 figs. (two males verso, one is a misidentification); Garwood & Lehman, 2011: Garwood & Lehman, 2011: 280 figs. (five figures misidentifications, from Guatemala and Honduras); Marín, 2011: 49 fig. 71, 56, 59, 66, 69, 71 (misiden- tification)]
Satyrotaygetis incerta (Butler & Druce, 1872),
Taygetis incerta Butler & Druce, 1872: 98. [TL: Costa Rica]
Taygetis incerta Butler & Druce; Butler, 1873: 149, pl.
53, fig. 1; Butler & Druce, 1874: 335; Kirby, 1877:
711; Godman & Salvin, 1880: 85; Lamas, 2004: 222 (last two as a synonym of S. satyrina (H. W. Bates)) Euptychia satyrina (H. W. Bates) f. incerta (Butler &
Druce); Weymer, 1911: 212.
Euptychia satyrina (H. W. Bates) var. incerta (Butler & Druce); Gaede, 1931: 464.
[Euptychia satyrina (H. W. Bates); Hayward, [1964]: 513 fig. 101, 514 (misidentification)]
Satyrotaygetis incerta (Butler & Druce); Forster, 1964: 73, t. 29, fig. 5.
[Cissia satyrina (W. H. Bates); DeVries, 1987: 275, pl.
48, figs. 8 (male), 9 (male) (misidentification)] [Euptychia satyrina H. W. Bates; d’Abrera, 1988: 781,
row [4], fig. [2] (male dorsal, pale form) (misidenti- fication)]
[Satyrotaygetis satyrina (H. W. Bates); Beccaloni et al.
2008: 343 (misidentification)]
Satyrotaygetis satyrina (H. W. Bates, 1865)
Taygetis satyrina H. W. Bates, 1865: 179. [TL: Guate- mala]
Taygetis satyrina H. W. Bates; Butler, 1868: 13; Kir- by, 1871: 110; Butler & Druce, 1874: 335; Marín,
2011: 63.
Euptychia satyrina (H. W. Bates); Butler, 1877: 120; Godman & Salvin, 1880: 84, pl. 8, fig. 23; Godman,
1901: 653 (misidentifications in part); Weymer,
1911: 212, pl. 48, row e; Gaede, 1931: 464; Ha-
yward, [1964]: 513 fig. 1001 (male genitalia), 514 [misidentification of S. incerta (Butler & Druce)]; d’Abrera, 1988: 781, row [4], figs. [1] (male dorsal, dark form), [2] (male dorsal, pale form, misidentifi- cation of T. incerta Butler & Druce), row [5] (male
M.G. Andrade-C., A.L. Viloria, E.R. Henao-Bañol & J.-F. Le Crom
ventral, pale form, misidentification of E. gigas But- ler), 789, 877.
Satyrotaygetis satyrina (H. W. Bates); Forster, 1964: 73, 75 fig. 34 (venation), t. 29, fig. 6; Lamas, 2004:
222; Murray & Prowell, 2005: 70, 72, 73, 75, 76, 77;
Chacón & Montero, 2007: pls. 177 figs. (male ver- so, one is a misidentification of S. gigas Btlr.), 178 fig. (pupa); Glassberg, 2007: 140, row [2], fig. [2];
Peña et al., 2010: 248, 250 fig. 2, 251 fig. 3, 253 fig.
4; Garwood & Lehman, 2011: 280 figs. (only one specimen correct [western Chiriqui, Panama]; five misidentifications of S. gigas (Butler)[Guatemala and Honduras])
[Cissia satyrina (W. H. Bates); DeVries, 1987: 275, pl.
48, figs. 8 (male), 9 (male) (misidentification of S. incerta (Butler & Druce))]
Cissia satyrina (W. H. Bates); Shou et al., 2006: 83. [Satyrotaygetis satyrina (H. W. Bates); Beccaloni et al.,
2008: 343 (misidentification of S. incerta (Butler & Druce), Marín, 2011: 49 fig. 71, 56, 59, 66, 69, 71
(misidentification of S. gigas (Butler))]
Satyrotaygetis tiessa (Hewitson, 1869), comb. nov. Euptychia tiessa Hewitson, 1869: 37. [TL: Ecuador] Euptychia tiessa Hewitson; Butler, 1870: 48, pl. 18, fig.
4; 1877: 120; Kirby, 1871: 643; 1879: 135; Butler
& Druce, 1874: 336; Godman & Salvin, 1880: 84;
Weymer, 1911: 212; Gaede, 1931: 466; d’Abrera,
1988: 764, 765, row [4], figs. [1] (male dorsal), [2]
(female ventral), row [5], fig. [1] (male ventral), 877.
Cissia tiessa (Hewitson); DeVries, 1987: 275, pl. 48, fig.
10 (male); T. & L. Racheli, 2001: 329; Shou et al., 2006: 83.
Magneuptychia tiessa (Hewitson); Lamas, 2004: 220;
Piñas, 2004: 34, figs. 263, 264; Murray & Prowell,
2005: 69, 72, 73, 75, 76, 77; Valencia-Martínez et al.,
2005: 167, pl. 33 fig. 3 [misidentification of Mag- neuptychia libye (Linnaeus)]; Chacón & Montero, 2007: pl. 175 fig. (male verso); Orozco et al., 2009: 37; Salinas et al., 2009: 436; Garwood & Lehman, 2011: 273, figs. (female and male), 80, fig.; Vargas & Salazar, 2014: 273.
E.[uptychia] tiessa Hewitson; Costa et al., 2016: 199. [¿?] tiessa Hewitson; Costa et al., 2016: 204.
Material examined
Satyrotaygetis gigas: MEXICO: 1 female, Oajaco [sic],
pur. From Hartwegg 44-13, Mexico, Type, B. M. Type No. Rh 3245, Euptychia gigas [female] Butler [Syntype] (NHMUK); HONDURAS: 1 male, SMD (STMD-D).
Satyrotaygetis incerta: COSTA RICA: 1 male, Costa
Rica, Van Patten, Druce Coll. Taygetis incerta Butl. & Dr. Type. Godman-Salvin Coll. 1904-1, B. C. A. Lep. Rhop. Euptychia satyrina Bates, B. M. Type No. Rh. 3246, Eupty- chia incerta [male] Butl. & Dr. T. incerta Butl. Type. Type
H. T. [Holotype] (NHMUK); PANAMA: 1 male SMD
(SMTD-D); 2 female, SMD, (SMTD-D). Satyrotaygetis
satyrina: GUATEMALA: 1 male, Centr. Valleys, Guate-
mala, F. D. G. & O. S. Type, Sp. figured, Godman-Salvin Coll. 1904-1, B. C. A. Lep. Rhop. Euptychia satyrina Bates, B. M. Type No. Rh. 3244, Euptychia satyrina [male] Bates, Type, H. T. [Holotype] (NHMUK). Satyrotaygetis tiessa: COLOMBIA: Chocó: 1 male, El Tabor, Chocó, 1500 m, 10.v.[19]90, leg. J. Salazar; 1 male, same data, 1800 m; 1 male, San José del Palmar, 1800 m, viii.[19]94, leg. D. Acosta; Cauca: 1 male, Tambito, 1000 m, 23.iii. [19]96, leg. T. Pyrcz (Genit. JFL 619 [fig. 21]) ( JFLC);
Montero ( JFLC); 1 male, Mistrato, Alto de Pisones, 1725 m, 3.vi.1992, leg. G. Andrade-C., GAC 3461, ICN-
MHN-L 8983; 1 male, same data, 1740 m, 4.vi.1992, leg.
Andrade-C., GAC 3463, ICN.MHN-L 8985 [Wing
prep. ALV053-17 (fig. 7), Genit. prep. ALV569-17]; 3 fe- males, Mistrato, San Antonio del Chami, camino al alto de Pisones, 1500, 17.iv.1993, leg., G. Andrade-C., GAC 4609, 4615, 4606, Gen. No. 1977, ICN-MHN-L 28535,
11295, 28538; 1 male, Santa Cecilia, x.1982, leg. J. Vélez, MHN-UC 12 (MHNUC); Valle del Cauca: 1 male, 1
female, Queremal, km 55, 1200 m, 10. iv. [19]90, leg. J. Salazar ( JFLC); 1 male, Queremal, Kilómetro 55, 278 m, 20.xii.1984, leg. J. Salazar, MHN-UC 278 (MHNUC); 1
male, Dagua, Columb occ. 95 Kalbr, SMD Museum Ber- lin. ECUADOR: 1 male, Ecuador, Hewitson Coll. 79-69, Euptychia tiessa Hew. 2., B. M. Type No. Rh. 3150, Eup- tychia tiessa [male] Hew., Type [Syntype] (NHMUK); Esmeraldas: 2 males, Esmeraldas, 500 m, viii.1959, leg. L. Richter, ICN-MHN-L 2698, 5929; 2 females, same data, vii.1959, leg. L. Richter, ICN-MHN-L 2709, 2703.
DeVries (1987: 275) provided some information on the natural history of S. tiessa. Individuals referred as to S. sa- tyrina from Costa Rica are certainly representatives of S. incerta (Butler & Druce), sp. restit. To our knowledge, S. satyrina, described from Guatemala, is distributed only in this country and the southern part of Mexico.
Nota bene: as a complement to the study of Magneup- tychia included in Costa et al. (2016) we present here three cases of misidentification of Magneuptychia lethra (Möschler) as Magneuptychia libye (Linnaeus) (figs. 8 wing venation, 16 male genitalia), type species of its genus: Bar- cant, 1970: pl. 13, fig. 17; Lewis, 1973: pl. 60, fig. 2 [same
Generic reassignments of satyrine butterflies
error in edition of 1987]; Urich, 1978: 12, figs., 13 [error pointed out by Cock (2014: 12), but the latter used the name M. newtoni (Hall), a junior synonym of M. lethra].
CONCLUSIONS
A number of independent phylogenetic studies based on the comparative analyses of DNA sequences, both mitochondrial (COI, Cyt b and ND1) and nuclear (EF- 1α, wingless, GAPDH y RpS5), have revealed paraphyly and polyphyly among many of the taxa conveniently ac- commodated under Magneuptychia Forster by most au- thors (Murray 2001, Murray & Prowell 2005, Peña et al. 2006, 2010, 2011; Marín et al. 2009, 2011, 2012, 2017;
Marín 2011). For instance, Murray (2001) and Murray
& Prowell (2005) included in their analysis six species of “Magneuptychia” and none formed monophyletic groups with any other (Murray, 2001: 360; Murray & Prowell, 2005:72). In all variants of their cladograms “Magneup- tychia” moderata appeared, well supported, as the clos- est taxon to Splendeuptychia ashna (type species of the genus). There was also strong support for the closest ge- netic distance between M. tiessa and S. satyrina (Murray 2001: 282, fig. 4.2; Murray & Prowell 2005: 77). Peña et
al. (2010: 247, 250 fig. 2, 251 fig. 3, 253 fig. 4) got Sa- tyrotaygetis satyrina notably separated (and nested among other genera) from any “Magneuptychia” included in their analyses, and “Magneuptychia” moderata and “Splendeup- tychia” purusana closely associated as sister taxa in the last branch of every cladogram (Figs. 2, 3, 4). This collection of evidence is congruent with the taxonomic hypotheses herein proposed for Nubila, gen. nov., and Satyrotaygetis, gen. rev. Not only it is not in conflict with our generic ar- rangement, but in fact it appears to support it.
According to our morphological observations Nubila, gen. nov., should be considered one sister genus to Splen- deuptychia, and quite apart from Magneuptychia.
Based on a morphology study in which a considerable number of characters were coded, Marín (2011) generated a cladogram in which “Magneuptychia” inani emerged very distant (sister to the Pareuptychia clade, pp. 56) from Magneuptychia libye (type species of the genus, derived in an unnamed group close to the Megisto clade), and at least once very close to Satyrotaygetis satyrina (pp. 59). On the other hand Murray (2001) studied, described and com- pared some morphological characters of the early stages of
M. libye (later published by Kaminsky & Freitas, in 2008) and “Magneuptychia” ayaya (a junior synonym of Macro- cissia iris, comb. nov.). She found considerable differences between the two taxa. These cases account for additional elements to justify the erection of Macrocissia, gen. nov.
It is morphologic and genetically distinct, but comparable and very close to the genus Satyrotaygetis (see Marín et al. 2017).
Colombeia, gen. nov. and Omacha, gen. nov. are rather peculiar, so far monospecific genera, whose relationships with other equivalent species groups of the Satyrinae can- not yet be established. Probably Colombeia, gen. nov. is truly related to Magneuptychia.
ACKNOWLEDGEMENTS
We express our gratitude to G. Lamas (Lima, Peru),
S. Attal (Paris, France), J. Diller (Münich, Germany),
O. Mielke (Curitiba, Brazil), L. D. Miller (†) (Sarasota, USA), G. Onore (Quito, Ecuador), T. Racheli (Roma, Italy) for providing bibliographic resources along the years. J. Camacho, I. Dorado, E. Inciarte and M. Quiróz (†) kindly assisted ALV during several visits to MALUZ to study satyrine butterflies. Q. Arias, J. Clavijo, J. De- Marmels, M. Gaiani and F. Romero patiently allowed us for extensive searches through the drawers of the MIZA and helped ALV as much as possible in locating disperse interesting material in the MIZA collections. We thank both Venezuelan institutions for their generous loan of specimens. ALV thanks M. Costa (Caracas) for his in- spiring enthusiasm and cooperation along the years, and P. R. Ackery, B. D’Abrera (†), J. Reynolds and R. I. Vane-Wright (London) for granting him open access to the collections and providing much professional help at the NHMUK, at least twenty years ago. Wilmer Rojas and Héctor Suárez (IVIC) assisted ALV in producing the illustrations of this article. Thanks to Julián Salazar (Museo de Historia Natural, Universidad de Caldas, Manizales), Fernando Trujillo (Scientific Director) and Federico Mosquera of the Omacha Foundation, and to Luis Germán Naranjo, Director of Conservation of WWF Colombia. The latter invited MGA to the field and funded fieldwork and information gathering for the declaration of the Tarapoto lakes as Ramsar site (now de- clared). MGA and ERHB received special support and funding from ICN during several phases of this research. MGA also wants to acknowledge his wife Angela Suárez and daughters Sofia Andrade Suárez and Elisa Andrade Suárez for their company and support in fieldwork at Puerto Nariño, Amazonas. MGA especially thanks
W. Mey (MNKB), A. Hausmann (ZSM) M. Nuß, A. Hundsdoerfer, Ch. Schmidt, F. Petzhold and U. Kall- weit (SMTD) for access to the collections under their care and for their multiple support during his recent sabbatical stay in Germany. ERHB thanks the SINCHI Institute, especially Dairon Cárdenas and Mariela Os-
M.G. Andrade-C., A.L. Viloria, E.R. Henao-Bañol & J.-F. Le Crom
orno, together with the Colombia Bio Expedition team for their support and collaboration. This article has im- proved by the criticism and suggestions of John Lynch, Gilson Rivas and a third, anonymous reviewer.
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Genus
Upperside Underside
Forewing Hindwing Forewing Hindwing
Magneuptychia Lacks ocelli Lacks ocelli
Ocelli not well developed, except for subapical, bipupillated
Ocellar series: R5-Cu1+A2 (obsolete), generally bipupillated but sometimes fused as one
Huberonympha Lacks ocelli Only females with
blind, subanal ocellus
With single subapical ocellus, bipupillated. Other vestigial on M1
Ocellar series: R5-Cu1, all five bipupillated (white)
Cissia With one subapical ocellus, bipupillated
Stevenaria Lacks ocelli
Only females with
With subanal ocellus, bipupillated
With subapical dark spot (M1) and a very diffuse dark vertigial spot on Cu1
With submarginal ocellar element on Cu1
With single subapical (M1) ocellus, bipupillated
With vestigial ocelli
Subapical ocellus on M1; vestigial ocelli
Ocellar series: R5-Cu1. R5, M2 and M3 only double spots (pupillae and spots all silver)
Ocellar series: R5-Cu1. R5, M2 and M3 without black; all five bipupillated (R5, M2, M3 silver; M1 Cu1 white)
Ocelli prominent on R5, M1 and
Satyrotaygetis
diffuse ocellar dark mark (subapical)
(monopupillated),
vestigial or smaller
in males. Better
developed in females
may variably appear on R5, M2 and M3. Monopupillated
notably Cu1. Absent, vestigial or
poorly developed (females) on M2
and M3. All monopupillated (white)
Colombeia
Only females with subapical dark mark (M1), and one dark vestigial on M3
Females with subanal blind ocellus (Cu1), with diffuse yellow ring; males only
dark spot, smaller
Ocellar series M1-M3 (Cu1 in females). All bipupillated (white); M1 prominent, subapical: M2
fused to M1
Ocellar series R5-M1, M3-Cu1 (M2 generally missing in males, vestigial in females). M3 and Cu1 fused. All bipupillated (white)
Macrocissia Lacks ocelli
Nubila Lacks ocelli
May exhibit dark ocellar element (Cu1), specially females
May exhibit two diffuse dark ocellar spots (M1, Cu1)
Subapical monopupillated ocellus (M1)
Lacks ocellar elements
Ocellar series R5-Cu1. R5 and M1bipupillated; Cu1 either with one or two pupillae, always exceeding limits of cell; M2
and M3 with double pupilla, elongated, silvery; lacking black
Ocellar series R5-Cu2. R5, M1, Cu1 and Cu2 fully ocellated (one pupil, white), M3,M3 lacking black, elongated, double pupillae, silver
Omacha Lacks ocelli Single, monopupillated ocellus (Cu1)
Exhibit one diffuse dark ocellar spot on
With vestigial subapical ocelli
Ocellar series R5-Cu1 complete.
All ocelli monopupillated
Ocellar series R5-Cu1 complete. R5 and M1 monopupillated, M2
Splendeuptychia Lacks ocelli
Cu1. Sometimes
one even less
distinct on M1
Lacks ocellar elements
to Cu1 bipupillated. Obsolescent or vestigial ocellar mark on Cu2
