Research Article |
Ryan A. St. Laurent‡, Jason J. Dombroskie‡
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Corresponding author: Ryan A. St. Laurent ( rstlaurent@flmnh.ufl.edu ) Academic editor: Alberto Zilli
© 2016 Ryan A. St. Laurent, Jason J. Dombroskie.
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:
St Laurent RA, Dombroskie JJ (2016) Revision of the genus Menevia Schaus, 1928 (Lepidoptera, Mimallonoidea, Mimallonidae) with the description of 11 new species. ZooKeys 566: 31-116. https://doi.org/10.3897/zookeys.566.6982
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Abstract
The Neotropical genus Menevia Schaus, 1928 is revised to include 18 species, 11 of which are new. Two species, M. ostia comb. n. and M. parostia comb. n. are transferred from Pamea Walker, 1855 to Menevia. Four species-groups are diagnosed for the first time based on external characters and male genitalia morphology. The following new species are described: M. rosea sp. n., M. torvamessoria sp. n., M. magna sp. n., M. menapia sp. n., M. mielkei sp. n., M. australis sp. n., M. vulgaris sp. n., M. franclemonti sp. n., M. vulgaricula sp. n., M. cordillera sp. n., and M. delphinus sp. n.. A neotype is designated for Mimallo plagiata Walker, 1855, which has since been placed in Menevia. Mimallo saturata Walker, 1855 is interpreted to be a nomen dubium.
Keywords
Mimallo , Neotropical, neotype, Pamea
Introduction
Until recently, very little revisionary work has been done with the family Mimallonidae.
Adding to the poor understanding of the higher-level arrangement of the family are a few weakly diagnosed genera currently persisting as catch-alls to subsume numerous recently described species (
The genus Menevia currently consists of five Central and South American species: M. plagiata (Walker, 1855), M. lantona (Schaus, 1905), M. lucara (Schaus, 1905), M. alurca Herbin & Mielke, 2014, and M. pallida Herbin & Mielke, 2014. No synonyms or other names have been formerly assigned to Menevia. We diagnose the genus Menevia based on external characters, re-describe all five currently known species, describe each previously undescribed female, describe 11 additional new species, and move two species currently assigned to the genus Pamea Walker, 1855 into Menevia.
Methods
Dissections were performed as described by
Costa Rican material of M. ostia was obtained from the biodiversity inventory of the Área de Conservación Guanacaste (ACG) (
The holotypes and one neotype were all dissected or, when present, previously made genitalia preparations were examined.
Specimens from the following collections were examined:
CGCM Collection of Carlos G. C. Mielke, Curitiba, Paraná, Brazil
CMNH
MEM Mississippi Entomological Museum, Mississippi State, Mississippi, USA
MGCL
McGuire Center for
MNHU
OM Collection of Olaf Hermann Hendrik Mielke, Curitiba, Paraná, Brazil
RAS Research collection of Ryan A. St. Laurent, Ithaca, New York, USA
The symbol ‡ will be used to represent unavailable names in the text (
Figures were manipulated with Adobe Photoshop CS4 (Adobe 2008). Male genitalia are figured in natural color with CS4 “auto color” used to improve white backgrounds. Female genitalia were treated with “auto tone” in CS4 to darken characters; insets, however, are manipulated only with “auto color.” Most adult specimens were photographed in natural light with an Apple iPhone 5S, additional adult photos were provided by CGCM and
Results and discussion
Menevia
Type species
Cicinnus lantona Schaus, 1905;
Diagnosis
Menevia can be recognized by the contrast between the usually gray submarginal area and the darker gray, brown, yellowish, or rarely pink medial area of the forewing and the presence of a white apical dash. This apical dash becomes the “postmedial lunule” (see Fig.
Description
Male.Head: Small, scales on frons swept ventrad, either the same color as vertex or darker ventrally, eyes large comprising about half to two-thirds of head area, eyes usually bordered posteriorly by darker collar of scales reaching labial palpi, labial palpi small, segments variably defined ventrally depending on thickness of vestiture, incrementally smaller in length distally, dorsally and laterally with darker scales contrasting with overall lighter coloration of head. Antenna bipectinate to tip, scape and pedicel tufted. Ocelli and chaetosemata absent. Thorax: Tan, gray, or straw colored. Densely covered in scales of varying widths with interspersed darker petiolate scales, scales of prothoracic collar finer, lighter, overlapping scales of mesothorax. Legs: Vestiture thick, scales long, especially on femur and tibia, coloration as for thorax, petiolate scales present. Tibial spurs often scaled, about one fourth length of tibia, apex may be somewhat hooked. Forewing dorsum: Forewing length: 14–28 mm. Triangular, outer margins concave on apical half, apex usually falcate. Ground color yellow, brown, gray, or gray-brown, lightly or moderately speckled by dark petiolate scales. Discal spot absent or faintly marked by light gray, no hyaline patches present. Dark postmedial line always present, either straight or with slight undulations. Gray submarginal area usually contrasting with medial area, submarginal area with a variably distinct white line originating from apical dash, white mark follows postmedial line from apex to one quarter to one half the length of postmedial line until sweeping outward toward wing margin, either at an acute angle or nearly perpendicular to postmedial line. In some species, white line wider, forming a complete or interrupted band following postmedial line, not sweeping toward wing margin. Antemedial line, if present, faint and undulated. Forewing venter: As in forewing dorsum but postmedial line may be fainter, antemedial line absent, discal spots may be much darker. Hindwing dorsum: Rounded or subtriangular, anal angle often accentuated, similar coloration and patterning as forewings, vague postmedial lunules present but undulated or sharply zigzagged, never dramatically swept to wing margin, antemedial line absent. Hindwing venter: Following similar pattern as forewing venter, usually lighter, frenulum with single bristle. Wing venation: As for Cicinnus melsheimeri (Harris, 1841) in
Remarks
The genus Menevia consists of four clear species-groups, which are delineated here for the first time. The species-groups are readily differentiated by the adult morphology; namely the degree of sexual dimorphism, size, forewing maculation and shape, ground color, and male genitalia characteristics.
Key to species-groups of Menevia
| 1 | Forewing submarginal area with white, curved, lunule originating from apical dash (Fig. |
2 |
| – | Forewing postmedial lunule replaced by continuous or interrupted white band along exterior edge of postmedial line | plagiata species-group |
| 2 | Ground color light tan, yellow, or orange, contrasting with gray submarginal area, postmedial lunule usually very weakly curved toward wing margin, forewings weakly falcate | 3 |
| – | Ground color mixture of deep red-brown and dark gray, not strongly contrasting with gray submarginal area except for when medial area more red-brown than gray, postmedial lunule sharply curved toward wing margin, forewings falcate | lucara species-group |
| 3 | Sexual dimorphism pronounced. Gray submarginal area highly contrasting with yellowish or golden medial area, submarginal area always with small accessory white mark near tornus, hindwing postmedial line mostly straight. Female: forewings broad, nearly ovoid; male: phallus always with dorsal ridge or variously shaped protuberance | ostia species-group |
| – | Male and female hardly differentiable. Gray submarginal area moderately or sometimes only weakly contrasting with light tan, yellowish, orange, or rarely pinkish medial area, small white accessory mark near tornus absent in most species, hindwing postmedial line usually wavy, especially near anal margin. Male: phallus irregularly shaped, usually without dorsal ridge, but when present, anteriorly situated, triangular | lantona species-group |
lantona species-group
The lantona species-group, containing the type species M. lantona by original designation, is the only species-group with very weak sexual dimorphism. This group includes the following species: M. lantona, M. rosea sp. n., M. torvamessoria sp. n., and M. magna sp. n. The females of two species belonging to this group were available for study, and both are exceptionally reminiscent of the males, with only slightly more elongated forewings and meagerly larger size. Species belonging to the lantona species-group are generally small for the genus, containing some of the smallest species of Menevia. The shape of the forewings of both males and females are not overly elongated, barely falcate, with weakly curving to almost straight postmedial lunules, and with a light ground color, which is usually yellowish with light shades of brown and tan. One species however, is mostly pink. The flat, triangular, or cupped processes of the gnathos and the usually broad phallus characterize male genitalia of this species-group.
Key to lantona species-group
| 1 | Ground color orange, yellow-tan, or dark tan, never with any pink suffusion, dorsum of phallus variable | 2 |
| – | Ground color suffused with pink, phallus with well-defined dorsal protuberance | M. rosea sp. n. |
| 2 | Forewing postmedial line straight or convex, faint or very thick and contrasting, abdomen never with midventral stripe. Panama and northern South America | 3 |
| – | Forewing postmedial line with slight inward kink on inferior half, phallus with triangular dorsal ridge, venter of abdomen with quarter-length or complete midventral stripe. Southeastern Brazil | M. magna sp. n. |
| 3 | Ground color orange-yellow, forewing postmedial line very dark and contrasting, genitalia with tegumen nearly circular, uncus hooked, phallus pistol shaped and bent mesally, tubular and elongated distally | M. torvamessoria sp. n. |
| – | Ground color light tan, fading to yellow in older specimens, forewing postmedial line usually thin and not highly contrasting against surrounding color, tegumen rectangular or somewhat ovoid, especially when prominently constricted at base of gnathos, uncus barely hooked, phallus irregularly shaped but not bent mesally or tubular distally | M. lantona |
Menevia lantona
Cicinnus lantona Schaus, 1905: 327–328
Cicinnus lantona;
Cicinnus lantona;
Menevia lantona;
Menevia lantona;
Menevia lantona;
Menevia lantona;
Type material
Holotype, ♂: FRENCH GUIANA: St. Jean, Maroni, F. Guiana/ Collection Wm. Schaus/ Perophora lantona type Schaus/ Type No.: 8894 U.S.N.M./
Additional specimens examined
(65 ♂, 1 ♀ total) BRAZIL: Amazonas: 1 ♂, Reserva Ducke, km. 26 Manaus-Itacoatiara Highway: 19.V.1972, E.G., I. & E.A. Munroe, St. Laurent diss.: 3-9-15:1 (
Diagnosis
Sexual dimorphism is weak; the females are only slightly larger. Thus both sexes of M. lantona are recognizable by their small to moderate size, only slightly falcate forewing apices, and yellowish tan to gold ground color with gray highlights, especially near the discal region. The postmedial lunule is bright white and only barely curved outward toward the forewing margin, not sharply curving as in other species-groups. The phallus is broad, usually with a small protuberance dorsally but always without a dorsal ridge. The gnathos processes are unique in that they are very broad, flat, and subtriangular, not oblong or thick as in most other species-groups. Female genitalia are separable from those of the similar female of M. magna sp. n. by the tergite of VIII, which is thin and upturned mesally rather than being replaced by a lightly sclerotized sac as in M. magna sp. n.
Description
Male.Head: Straw or tan colored, eyes bordered posteriorly by dark brown collar of scales reaching labial palpi, labial palpi small, segments weakly defined ventrally due to ventral tufts, dorsally with darker scales contrasting with overall straw coloration. Scape and pedicel weakly tufted. Thorax: As for genus but tan or gold, fading to straw. Legs: As for genus. Tibial spurs thin apically, terminal third not scaled, especially ventrally. Forewing dorsum: Forewing length: 15–18 mm, avg.: 16.2 mm, n = 28. Triangular, apical half of outer margins concave, convex near tornus, apex slightly falcate. Ground color yellowish tan to gold with varying degrees of gray, especially near discal region, overall lightly speckled by dark petiolate scales. Discal spot faintly marked by light gray. Apex marked by black scales above apical dash. Straight or slightly undulating postmedial line black or brown. Antemedial area lighter, submarginal area gray with tan coloration near tornus, postmedial lunule originating from near where apical dash meets postmedial line, lunule follows postmedial line from apex to one third length of postmedial line where lunule smoothly curves outward toward wing margin, forming roughly 45 degree angle with postmedial line. Antemedial line very faint or absent, if present, brown, undulating. Forewing venter: As in forewing dorsum but postmedial line fainter, antemedial line absent; usually rounded discal spot present, small, black. Hindwing dorsum: Rounded with margin weakly pointed mesally, anal angle weakly accentuated, similar coloration and patterning as forewings, vague postmedial lunule originating near anterior margin undulating, not steeply swept to margin, antemedial line absent, postmedial line undulating, especially near anal angle. Hindwing venter: Following similar pattern as forewing venter but discal mark not always present. Abdomen: As for genus. Coloration a continuation of tan or golden thoracic color. Midventral stripe absent. Genitalia: (Fig.
Distribution
Remarks
As a wide-ranging species, M. lantona expresses some geographic variation. Most notably, specimens from Colombia are larger (Fig.
Menevia lantona species-group adults, a recto, b verso. 6 M. lantona holotype ♂, French Guiana, St. Jean du Maroni (
Menevia rosea sp. n.
Type material
Holotype, ♂: ECUADOR: Ecuador: Napo, Simon Bolivar [Simón Bolívar], Coca River canyon, 1200 m, 16 Aug 1996, Jan Hillman, undisturbed wet forest/ St. Laurent diss.: 3-7-15:9/ HOLOTYPE male Menevia rosea St Laurent and Dombroskie, 2016 [handwritten red label]/ (CMNH). No paratypes. Type locality: Ecuador: Napo: Simón Bolívar.
Diagnosis
Menevia rosea is distinguishable from all other species in the lantona species-group by the pink coloration of the forewings, especially medially and proximal to the apical region. The postmedial lunule is also more weakly curved than in the most similar species, M. lantona. Genitalia characters should also readily distinguish M. rosea from other species in the lantona species-group. The particularly short phallus has a small, but obvious dorsal ridge, which is lacking in M. lantona. The phallic ridge of M. rosea should not be confused with the dorsal protuberance on the phallus of some M. lantona, as this protuberance is closer to mid-length of the phallus while the phallic ridge of M. rosea is more distal. The juxtal processes are more curved toward the distal end of the phallus. The valves are more triangular and are particularly broad proximal to the vinculum, the saccular edges of the valves are also particularly straight.
Description
Male.Head: Brownish gray with pinkish hue, eyes bordered posteriorly by dark brown collar of scales reaching labial palpi, labial palpi small, segments weakly defined ventrally due to ventral tufts, dorsally with darker scales contrasting with overall pinkish gray coloration. Scape and pedicel weakly tufted. Thorax: As for genus but light tan, scales of prothoracic collar pinker, tipped with white. Legs: As for genus. Tibial spurs very thin, relatively long, terminal third not scaled, especially ventrally. Forewing dorsum: Forewing length: 16 mm, n = 1. Triangular, apical half of outer margins concave, apex slightly falcate. Ground color light tan with excessive pink scaling, especially medially and nearing apex before postmedial line, very sparsely speckled by dark petiolate scales. Discal spot faintly marked by light gray. Apex marked by black scales above small apical dash. Dark brown postmedial line mostly straight, somewhat undulating. Antemedial area lighter, less pink, submarginal area faint gray to more tan near tornus, postmedial lunule originating perpendicular to where apical dash meets postmedial line, lunule follows postmedial line from apex to nearly half length of postmedial line where lunule barely curves outward toward wing margin, forming very acute angle with postmedial line. Antemedial line absent. Forewing venter: As in forewing dorsum, pink coloration widespread but postmedial line fainter, more curved, antemedial line absent, small black discal spot present. Hindwing dorsum: Rounded with margin weakly pointed mesally, anal angle very weakly accentuated, similar coloration and patterning as forewings but with more petiolate scales, vague postmedial lunule originating near anterior margin undulating, not steeply swept to margin, antemedial line absent, postmedial line mostly straight, brown, surrounded by pink. Hindwing venter: Following similar pattern as forewing venter, but lighter. Abdomen: As for genus. Coloration a continuation of tan coloration of thorax with pink hue. Genitalia: (Fig.
Distribution
(Map
Etymology
Menevia rosea is named for the unique pink coloration of this species, unlike other species of the lantona species-group, which are usually tan.
Remarks
This new species is unique externally and is the only species in the lantona species-group with pink scales on the wings. Pink scaling is seen in many other Menevia species, particularly those in the lucara species-group. However, all other characteristics of patterning and the genitalia of M. rosea perfectly match those characters typical of the lantona species-group.
Apart from the interesting external coloration and genitalia characters, the type locality of this species is noteworthy mostly due to the relatively high elevation, 1200 m. The most similar species, M. lantona, has been collected in the Napo province just 80 km south of the type locality of M. rosea. Furthermore, the specimen collected 80 km south of the M. rosea type locality was collected at 1097 m elevation, also high for the genus. We have attributed this particular specimen to M. lantona due to the complete lack of pink and the genitalia characters. It is worth noting however, that this specimen has much darker postmedial lines on the fore and hindwings than typical M. lantona. Although the external characters and elevation of the collecting site are somewhat unique, the genitalia characters (St. Laurent diss.: 7-7-15:3) are completely typical of M. lantona, and are unlike either M. rosea or the other suspected possibility, M. torvamessoria sp. n. described below. Other Ecuadorian M. lantona are from much lower altitudes, 380-400 m. The differences in genitalia and the fact that M. lantona has been recorded from rather high elevations suggest that M. rosea is not a mere high elevation form of M. lantona.
Menevia torvamessoria sp. n.
Type material
Holotype, ♂: PERU: La Union [La Unión], R. Huacamayo, Carabaya [Puno], 2000 ft., wet s., Nov. 1904 (G. Ockenden)/ Rothschild Bequest BM 1939–1/ St. Laurent diss.: 6-29-15:4/ BMNH(E) 1378762/ HOLOTYPE male Menevia torvamessoria St Laurent and Dombroskie, 2016 [handwritten red label] (
Paratypes, 2 ♂: BRAZIL: Pará: 1 ♂, Monte Cristo, Rio Tapajós: Dognin Collection,
Diagnosis
Externally, M. torvamessoria is similar to M. lantona, but can be distinguished by the darker, yellow-orange ground color (in well-preserved specimens); M. lantona is lighter, more yellow-tan. Additionally, the postmedial line is very dark and contrasting in M. torvamessoria and there is a roughly rectangular gray patch of scales that extends from the discal region to the postmedial line. There is a similar gray patch in M. lantona but is not so well defined. The most outstanding diagnostic features of this new species are in the male genitalia. The phallus is unlike any other in the genus, it is almost pistol shaped and sharply bent halfway along its length. The dorsal surface of the proximal end of the phallus bears a distinct triangular or rounded ridge while the remainder of the length of the phallus is smooth, elongated, and tubular. The juxtal processes are very thin, shorter and more curved in other species in the lantona species-group. The gnathos processes are unique in that they are cupped and circular. The valves are unlike the previous species in that they are symmetrical and do not bear teeth on the saccular edge, but instead have distinct sclerotized inward facing lobes at the base of the valves, which conceal the gnathos. The sides of the tegumen are greatly bowed outwards, causing it to appear almost circular. Finally, the scythe-like uncus is acutely triangular and sharply hooked.
Description
Male.Head: Brownish tan or almost black, eyes bordered posteriorly by dark brown collar of scales reaching labial palpi, labial palpi small, segments moderately well defined ventrally due to ventral tufts, dorsally with darker scales contrasting with overall lighter coloration. Scape and pedicel thinly tufted. Thorax: As for genus but light tan, fading to straw. Legs: As for genus. Tibial spurs relatively small, only lightly scaled, especially proximally. Forewing dorsum: Forewing length: 14–16.5 mm, avg.: 14.8 mm, n = 3. Triangular, apical half of outer margin concave, apex slightly falcate. Ground color orange-yellow with elongate, gray rectangle of scales extending from discal region to postmedial line, overall very sparsely speckled by dark petiolate scales. Discal spot faintly marked by light gray. Apex marked by black scales above scythe-like apical dash. Slightly undulating postmedial line black, strongly contrasting. Antemedial area lighter, submarginal area gray with slight invasion of medial area coloration near tornus, postmedial lunule originating from near where apical dash meets postmedial line, forming scythe-like dash, lunule follows postmedial line from apex to one third length of postmedial line where lunule smoothly curves outward toward wing margin, forming roughly 45 degree angle with postmedial line. Antemedial line very faint or absent, if present, brown, undulating, bowed out near anal margin. Forewing venter: As in forewing dorsum but postmedial line fainter, undulations more distinct, antemedial line absent, small black, rounded or oblong discal spot present. Hindwing dorsum: Rounded, anal angle weakly accentuated, similar coloration and patterning as forewings, but postmedial lunule almost nonexistent, antemedial line absent, postmedial line more undulated and brown, not black and contrasting, undulations prominent, especially near anal angle. Hindwing venter: Following similar pattern as forewing venter, but lighter, discal spot much less distinct or absent. Abdomen: As for genus. Coloration a continuation of tan thoracic color. Midventral stripe absent. Genitalia: (Fig.
Distribution
(Map
Etymology
Menevia torvamessoria is named for the unique hooked uncus, reminiscent of the Grim Reaper’s (=torva messor Latin) scythe. The name is doubly appropriate to describe the apical dash, which combined with the postmedial lunule of the forewing, is scythe-like, a character seen in all Menevia species.
Remarks
Although externally M. torvamessoria is rather similar to M. lantona, this new species is wholly unlike any other in the genus when taking into account male genitalia. Every aspect of the genitalia, particularly the hooked uncus, circular tegumen, cupped gnathos processes, bulbous projections at the base of the valves, and the shape of the phallus, are all unique to this species. Menevia torvamessoria belongs in the genus Menevia due to the presence of the general structure of the genitalia such as the paired gnathos, juxtal processes, and outward facing tusks; but it is difficult to assign this taxon to a species-group based on genitalia alone. External characters, however, such as the size, orange-yellow coloration, and weakly falcate forewings, tentatively allow placement of M. torvamessoria in the M. lantona species-group.
In addition to the unusual genitalia, the distribution is also strange, but this may be due to one specimen being incorrectly labeled. One paratype bears a nearly illegible label reading “Monte Cristo, Rio Tapajós, Amazonas” and seems to be from Monte Cristo, in the Brazilian state of Pará, on the Tapajós River. This particular location is very low in elevation, with some hills only as high as about 300 m nearby (as determined from Google Earth), which is quite divergent from the Andean foothill localities of the holotype and paratype. Either the specimen is mislabeled or M. torvamessoria is very widespread in South America, and apparently very rare. Regardless of the uncertainty of the collecting locality of this paratype, its genitalia display the very unique and bizarre characteristics of the specimens from southern Peru and the ground color of this specimen is the same distinctive orange-yellow of M. torvamessoria, thus we include this individual in the type series given that there are so few examples of this species available.
Menevia magna sp. n.
Type material
Holotype, ♂: BRAZIL: Santa Catarina: BRAZIL – SC, São Bento do Sul, Rio Natal, 450 m. I. Rank leg, 27.VI.2014, S 26°20'2", W 49°18'30" / St. Laurent diss.: 6-16-15:1/ DZ 32.694/ HOLOTYPE male Menevia magna St Laurent and Dombroskie, 2016 [handwritten red label]/ (
Paratypes, 22 ♂, 3 ♀: BRAZIL: Santa Catarina: 1 ♂, Blumenau: X, Pohl, “696”,
Diagnosis
Menevia magna is similar to other species in the lantona species-group, except M. rosea, due to the tan ground color, but can be easily recognized by the larger size on average, the darker gray and brown scaling overall (especially in fresh specimens), and the usually mesally kinked forewing postmedial line. Additionally, the male genitalia are unique in that the valves are very broad and ovoid, the paired processes of the gnathos very thin and fingerlike or acutely triangular, and the smoothly curved phallus with a prominent, sharp, triangular dorsal ridge. Female genitalia are unique in the genus in having a bulbous, sac-like, sclerotized structure dorsally, replacing the usual sclerotization of tergite VIII, which also distinguishes females of M. magna from the very similar female of M. lantona.
Description
Male.Head: Tan to gray, eyes bordered posteriorly by dark brown collar of scales reaching labial palpi, labial palpi small, segments weakly defined ventrally due to ventral tufts, dorsally with very dark scales contrasting with overall tan-gray coloration. Scape and pedicel weakly tufted. Thorax: As for genus, but tan, fading to straw. Legs: As for genus. Tibial spurs short, mostly scaled. Forewing dorsum: Forewing length: 17–19.5 mm, avg.: 17.8 mm, n = 6. Triangular, apical half of outer margin concave, apex falcate. Ground color dark tan-brown fading to lighter yellowish, grayer medially, overall lightly speckled with dark petiolate scales. Discal spot faintly marked by light gray. Apex marked by black scales above apical dash. Dark brown postmedial line straight with slight inward kink mesally. Antemedial area lighter, submarginal area gray with very little tan coloration near tornus, postmedial lunule originating from near where apical dash meets postmedial line, lunule follows postmedial line from apex to one third length of postmedial line where lunule angled outward toward wing margin, forming roughly 45 degree angle with postmedial line. Antemedial line usually present, brown, undulated. Forewing venter: As in forewing dorsum but postmedial nearly absent, may be more heavily speckled with petiolate scales, antemedial line absent, small black, rounded or oblong discal spot present. Hindwing dorsum: Rounded with margin weakly pointed mesally, anal angle accentuated, similar coloration and patterning as forewings, vague postmedial lunules originating near anterior margin undulating or nearly straight, antemedial line absent, postmedial line undulating, especially near anal angle. Hindwing venter: Following similar pattern as forewing venter but lighter, discal spot absent. Abdomen: As for genus. Coloration a continuation of tan thoracic color. Midventral stripe weak, either extending from posterior tip to about one quarter length of abdomen or until thorax. Genitalia: (Fig.
Distribution
(Map
Etymology
Menevia magna is named for the very broad valves of the male genitalia and also for the overall size, which is quite large on average for the lantona species-group.
Remarks
Menevia magna appears to be closely related to the much more widespread M. lantona of northern and central South America. Based on our present understanding of the distribution of this species in São Paulo, Santa Catarina, and Rio de Janeiro, it can be reasonably inferred that M. magna is probably found in eastern Paraná as well.
lucara species-group
Species of the lucara species-group are similar to those of the lantona species-group and includes M. lucara, M. menapia sp. n., and M. mielkei sp. n. Sexual dimorphism is more pronounced than in the previous group, but still not as strong as in the next two species-groups, with females having broader, more ovoid forewings than males. The size of the species belonging to this group are on the smaller side for the genus, being only slightly larger than those of the lantona group overall. Forewing shape is stout and triangular as in the lantona group, but with more acutely falcate forewings and more steeply curved postmedial lunules. The ground color is dark, generally slate gray with varying degrees of brown and red in the medial area. The lobes of the gnathos in this species-group are unique among the genus, rather than being flattened, cupped, triangular or subtriangular in shape, they are thick, usually upturned, well-sclerotized structures reminiscent of boxing gloves. The phallus is rather tubular and simple, with only one species having a noticeable dorsal phallic ridge.
Key to lucara species-group
| 1 | Forewing not acutely falcate, phallus without dorsal ridge. Central America to northern South America | 2 |
| – | Forewing very acutely falcate, narrow, phallus with dorsal ridge. Southeastern Brazil | M. mielkei sp. n. |
| 2 | Juxtal processes pointed. Belize and Guatemala | M. menapia sp. n. |
| – | Juxtal processes terminally spatulate. Northern South America | M. lucara |
Menevia lucara
Cicinnus lucaraSchaus, 1905: 328
Menevia lucara;
Menevia lucara;
Menevia lucara;
Menevia lucara;
Type material
Holotype, ♂: FRENCH GUIANA: St. Jean, Maroni, Fr. Guiana/ Collection Wm. Schaus/ Perophora lucara type Schaus/ Type No.: 8895 U.S.N.M./
Additional specimens examined
(60 ♂, 4 ♀ total) BRAZIL: Amazonas: 4 ♂, Hyutanahan [Huitanaã], Rio Purus: II.1922, III.1922, S.M. Klages, Carn. Mus. Accs. 6963, 7088, 8840, St. Laurent diss.: 2-5-15:5 (CMNH). 2 ♂, Nova Olinda, Rio Purus: V.1922, VI.1922, S.M. Klages, Carn. Mus. Acc. 7088, St. Laurent diss.: 2-5-15:9 (CMNH). 1 ♂, Miracema, Rio Purus: IV.1922, S.M. Klages, Carn. Mus. Acc. 6960, St. Laurent diss.: 2-7-15:1 (CMNH). 1 ♂, “en remontant l’Amazones de Teffé à Tonantins,” [between Tefé and Tonantins]: XI.1921, Dognin Collection,
Diagnosis
Both males and females are recognizable by their moderate size and gray ground color with pinkish highlights especially concentrated near the apical angle of the contrasting black postmedial line and near the anal angle of hindwings. The postmedial lunule is bright white and sharply curved outward toward the forewing margin, nearly forming a semicircle. The phallus/juxta combination is unique in the genus in that the phallus is smooth and cylindrical, without a dorsal phallic ridge. The terminal ends of the juxtal processes are spatulate, not pointed, and curved inward toward each other, near the terminal quarter of their length. The gnathos processes are highly distinct; they are thick, upturned, heavily sclerotized, somewhat boxing glove in shape.
Description
Male.Head: Light gray, straw colored in old specimens, eyes large comprising about two-thirds of head area, eyes bordered posteriorly by dark brown collar of scales reaching labial palpi, labial palpi small, segments weakly defined ventrally due to ventral tufts, dorsally with darker scales contrasting with overall gray coloration. Scape and pedicel tufted. Thorax: As for genus. Grayish. Legs: As for genus. Tibial spurs thin apically, terminal third not scaled, especially ventrally, weakly hooked. Forewing dorsum: Forewing length: 16.5–21 mm, avg.: 17.3 mm, n = 22. Triangular, apical half of outer margin concave, apex falcate. Ground color gray with pink hue especially near apical point of postmedial line and throughout medial area, overall lightly speckled by dark petiolate scales. Discal spot faintly marked by light gray. Apex marked by black scales above apical dash. Straight postmedial line black or brown, usually strongly contrasting. Antemedial area lighter, submarginal area gray without pink hue, somewhat contrasting with medial area, postmedial lunule originating from apical dash, lunule follows postmedial line from apex to one third length of postmedial line where lunule sharply sweeps outward toward wing margin, forming acute angle with postmedial line. Antemedial line usually absent, if present, faint, brown, undulating. Forewing venter: As in forewing dorsum but postmedial line fainter, antemedial line absent, small black discal spot present. Hindwing dorsum: Somewhat rounded with margin weakly pointed mesally, anal angle accentuated, similar coloration and patterning as forewings, vague postmedial lunule originating near anterior margin, sweeping outward to marginal point, antemedial line absent, postmedial line straight or slightly curved, especially near anterior wing margin. Hindwing venter: Following similar pattern as forewing venter, but slightly lighter, discal mark absent. Abdomen: As for genus. Coloration a continuation of grayish thoracic color. Midventral stripe absent. Genitalia: (Fig.
Distribution
Remarks
Like M. lantona, M. lucara is a wide-ranging Amazonian species found throughout northern South America and is sympatric with M. lantona throughout most of its range.
Some geographical variation has been noted among the material examined. Specimens from Panama and Colombia (Figs
Menevia lucara species-group adults, a recto, b verso. 18 M. lucara holotype ♂, French Guiana, St. Jean du Maroni (
Menevia menapia sp. n.
Type material
Holotype, ♂: GUATEMALA: Cayuga Guat/ June/ Schaus and Barnes coll/
Paratypes, 8 ♂: BELIZE: 2 ♂, Punta Gorda, Brit. Honduras: VI.1933, J.J. White, Rothschild Bequest BM 1939-1, St. Laurent diss.: 6-29-15:3, BMNH(E) 1378758 (
Diagnosis
Externally M. menapia is nearly identical to M. lucara, albeit this new species is slightly smaller on average. The most significant differences in external characters are present in the postmedial lunules of the fore- and hindwings. The postmedial lunule is less sharply swept to the forewing margin in M. menapia, and on the hindwing the lunule is more distinct, a brighter white rather than extremely faded as in M. lucara. The male genitalia of M. menapia are recognizable by the weaker sclerotization; smaller, slightly thinner processes of the gnathos, and the sharp, rather than spatulate, apices of the juxtal processes. Furthermore, the lobes at the base of the phallus are much shorter and stouter than in M. lucara. Geography is perhaps the easiest way to differentiate M. menapia from other species in the lucara species-group, as it is the only representative of the group from Central America north of Panama.
Description
Male.Head: Gray, fading to straw colored in old specimens, eyes large comprising about two-thirds of head area, eyes bordered posteriorly by dark brown collar of scales reaching labial palpi, labial palpi small, segments weakly defined ventrally. Scape and pedicel tufted. Thorax: As for genus. Gray fading to straw in old specimens. Legs: As for genus. Tibial spurs thin apically, terminal third not scaled, weakly hooked. Forewing dorsum: Forewing length: 15–17 mm, avg.: 16 mm, n = 7. Triangular, apical half of outer margin concave, apex falcate. Ground color gray with pink hue medially, overall lightly speckled by dark petiolate scales. Discal spot faintly marked by light gray. Apex marked by black scales above apical dash. Weakly concave postmedial line black, contrasting. Antemedial area lighter, submarginal area gray without pink hue, contrasting with pinkish medial area, postmedial lunule originating from apical dash, lunule follows postmedial line from apex to one third length of postmedial line where lunule sweeps outward toward wing margin, roughly forming 45 degree angle with postmedial line. Antemedial line faint or absent, if present, brown, undulating. Forewing venter: As in forewing dorsum but postmedial line fainter, antemedial line absent, small black discal spot present. Hindwing dorsum: Rounded with margin weakly pointed mesally, anal angle accentuated, similar coloration and patterning as forewings, postmedial lunule present, originating near anterior margin, sweeping outward and fading to marginal point, antemedial line absent, postmedial line straight except near anterior margin. Hindwing venter: Following similar pattern as forewing venter. Abdomen: As for genus. Coloration a continuation of grayish thoracic color. Midventral stripe absent. Genitalia: (Fig.
Distribution
(Map
Etymology
Menevia menapia is named for the likely derivation of Menevia, Menapia. The etymology of Menapia, however, is less clear and may refer to an ancient Roman settlement supposed to have existed in Pembrokeshire, Wales or to a settlement inhabited by the Menapii people in Belgica.
Remarks
This species, although very similar in external appearance and genitalia characteristics to the South American representatives of the lucara species-group, especially the Amazonian M. lucara, is separated by well over 2000 km land distance. Because of the extreme allopatry of M. menapia, it is surprising that this new species differs so little from the wide-ranging, South American M. lucara. Despite these similarities, the allopatry combined with the overall slightly smaller size, minor external differences, and the distinct genitalia differences, warrant the separation of these two similar species.
Menevia mielkei sp. n.
Type material
Holotype, ♂: BRAZIL: Minas Gerais: 29-I--3-II-2003, Estação Biológica de Caratinga, Caratinga, MG, 400 m, Mielke & Casagrande leg./ OM 61.563/ CGCM diss.: OM 61.563/ HOLOTYPE male Menevia mielkei St Laurent and Dombroskie, 2016 [handwritten red label] (donated to
Paratypes, 13 ♂, 1 ♀: BRAZIL: Espírito Santo: 1 ♂, Linhares, 40 m: 05–09.IV.1992, V.O. Becker col.,
Diagnosis
Externally M. mielkei is nearly identical to both M. lucara and M. menapia. The forewings of M. mielkei are slightly narrower than in the other two species, most notably by the somewhat more acute apices. Genitalia are very useful however, in the diagnosis of M. mielkei. In M. mielkei, the uncus is more slender, the processes of the gnathos are extremely atrophied and thin, the phallus has a distinct dorsal ridge, the processes of the juxta are nearly vertically extended above the base of the phallus, not curved over its length, and the valves lack the saccular edge tooth. Furthermore, the lobes at the base of the phallus are much shorter and stouter than in M. lucara, and in that way are more similar to M. menapia.
Description
Male.Head: Light gray, eyes large comprising about two-thirds of head area, eyes bordered posteriorly by somewhat reduced dark brown collar of scales reaching labial palpi, labial palpi small, segments weakly defined ventrally. Scape and pedicel tufted. Thorax: As for genus. Grayish. Legs: As for genus. Tibial spurs thin apically, terminal third not scaled, especially ventrally, weakly hooked. Forewing dorsum: Forewing length: 16–17 mm, avg.: 16.6 mm, n = 4. Triangular, apical half of outer margin deeply concave, slightly convex near tornus, apex acutely falcate. Ground color gray with pinkish-red to salmon hue especially near apical point of postmedial line and near discal region, overall lightly speckled by dark petiolate scales. Discal spot faintly marked by light gray. Apex marked by black scales above apical dash. Postmedial line straight, black-brown. Antemedial area lighter than medial area, submarginal area gray without pink hue, postmedial lunule originating from apical dash, lunule follows postmedial line from apex to roughly one third length of postmedial line where lunule sharply sweeps outwards toward wing margin, forming acute angle with postmedial line. Antemedial line very faint, brown, undulating. Forewing venter: As in forewing dorsum but postmedial line fainter, pinkish-red hue concentrated near costa and discal region, antemedial line absent, small black discal spot present. Hindwing dorsum: Rounded with margin weakly pointed mesally, anal angle accentuated, similar coloration and patterning as forewings, small black discal mark usually present, very vague postmedial lunules originating near anterior margin sweeping outward to marginal point, antemedial line absent, postmedial line straight or slightly concave. Hindwing venter: Following similar pattern as forewing venter, pinkish-red hue concentrated in anal region. Abdomen: As for genus. Coloration a continuation of grayish thoracic color. Midventral stripe absent. Genitalia: (Fig.
Distribution
Menevia mielkei is found in a small region of southeastern Brazil in the states of Rio de Janeiro, Espírito Santo, and adjacent eastern Minas Gerais. Whether or not this species is more widely distributed in the Brazilian Atlantic Forest is yet to be determined.
Etymology
Menevia mielkei is named after Carlos G. C. Mielke who provided extensive support and data throughout the process of writing this revision.
Remarks
Like M. menapia, M. mielkei is very similar to the wide-ranging M. lucara. Both M. menapia and M. mielkei are apparently restricted in distribution, both species being widely allopatric with M. lucara. Of the three species in the lucara species-group, M. mielkei has the most distinct genitalia characters as outlined in the diagnosis. The dorsal ridge of the phallus is a common character in many other species of Menevia, but within the lucara species-group, this ridge is only present in M. mielkei.
ostia species-group
The following group contains three species, M. ostia comb. n., M. parostia comb. n., and M. pallida. The former two species were assigned to the genus Pamea Walker, 1855 and the latter species was recently described in Menevia (
Key to ostia species-group
| 1 | Coloration washed out yellow-tan, hindwing with very faint zigzagged lunule; females: hindwing postmedial line closer to wing margin than to middle of wing | 2 |
| – | Coloration more gold yellow-orange, hindwing with usually prominent zigzagged lunule; females: hindwing postmedial line closer to thorax than to middle of wing. Distribution wide but usually in wet forests from Costa Rica to throughout Amazonia and into the Brazilian Atlantic Forest | M. ostia comb. n. |
| 2 | Female forewing less than 21 mm, male unknown | M. parostia comb. n. |
| – | Female forewing greater than or equal to 21 mm. Brazilian Cerrado, possibly also south to Paraguay | M. pallida |
Menevia ostia , comb. n.
Perophora ostia Druce, 1898: 447; Tab. LXXXVIII, fig. 18 ♀
Pamea ostia;
Pamea ostia;
Type material
Holotype, ♀: PANAMA: Chiriqui [Chiriquí]/ 855/ Coll. Staudinger/ Type/ Perophora ostia ♀ type Druce/ Coll. Staudinger II.1178./ St. Laurent diss.: 7-14-15:1/ (MNHU) [examined]. Type locality: Panama: Chiriquí.
Additional specimens examined
(31 ♂, 15 ♀ total) BRAZIL: Espírito Santo: 9 ♂, Linhares, 40 m: 20–29.II.1992, 06.III.1993, 25–30.I.1998, V.O. Becker Col., Col. Becker 80930, 82945, 113493,
Diagnosis
Menevia ostia can be differentiated from all other species-groups by the gold-yellow to pale yellow ground color with the contrasting silvery gray submarginal area with a small white accessory mark near the tornus. These traits combined with overall very weakly falcate forewings, distinguish M. ostia from the similar M. lantona but not necessarily from the similar M. pallida and M. parostia comb. n. to be diagnosed below. The width of the submarginal area of the hindwings of female M. ostia is wider than in both M. pallida and M. parostia females. The postmedial line is situated at about midway along the length of the hindwing in M. ostia, but is slightly closer to the wing margin in the other two species. Male genitalia are distinct, except when in comparison with M. pallida, in that the apical end of the phallus is sharply downturned and the dorsal ridge of the phallus is highly variable, though always present. In females, the lateral portion of the prominently sclerotized VIII has appendicular apophyses dorsolaterally in addition to the apophyeses anteriores, distinguishing the females from a questionable female of M. pallida, where the appendicular apophyses are absent.
Description
Male.Head: Straw or tan-gold colored, eyes bordered posteriorly by dark brown collar of scales reaching labial palpi, labial palpi small, segments somewhat well defined ventrally, dorsally with darker scales contrasting with overall lighter coloration. Scape and pedicel weakly tufted. Thorax: As for genus. Gold to pale, fading to straw. Legs: As for genus. Tibial spurs thin apically lightly to almost completely scaled. Forewing dorsum: Forewing length: 12.5–18.5 mm, avg.: 16.8 mm, n = 23. Triangular, apical half of outer margin weakly concave, apex slightly falcate. Ground color gold-yellow to pale yellow, overall very lightly speckled by dark petiolate scales. Discal spot faintly marked by silvery white. Apex marked by black scales above apical dash. Black postmedial line slightly concave, sometimes weakly undulating. Submarginal area silvery gray, postmedial lunule originating from near where apical dash meets postmedial line, white mark follows postmedial line from apex to one quarter length of postmedial line where mark smoothly curves outwards toward wing margin becoming diffuse, forming acute angle with postmedial line. White accessory mark present near tornus. Antemedial line very faint or absent, if present, brown, undulating. Forewing venter: As in forewing dorsum but postmedial line fainter, convex near tornus, antemedial line absent, small black, somewhat rounded or elongated discal spot present. Hindwing dorsum: Rounded, anal angle weakly accentuated, similar coloration and patterning as forewings, postmedial lunule vague or well defined, zigzagged, originating near anterior margin, following curvature of wing margin, not steeply swept to margin, antemedial area lighter, postmedial line straight, sometimes undulating near anal angle. Hindwing venter: Following similar pattern as forewing venter but lighter, discal mark absent, marginal area usually browner than surrounding area. Abdomen: As for genus. Coloration a continuation of thoracic color. Midventral stripe absent. Genitalia: (Figs
Distribution
(Map
Natural history
Menevia ostia is one of the few species of Menevia with known host plant associations. Rearing records from D. Janzen & W. Hallwachs show that M. ostia feeds on both Terminalia amazonia and T. oblonga (Combretaceae). Additionally, a female specimen from Suriname at the
Furthermore, D. Janzen provided us access to photos of reared M. ostia, published here for the first time (see Figs
Remarks
Druce described Menevia ostia at a time when most new Mimallonidae that were being described were being placed in the now invalid, preoccupied genus Perophora Harris, 1841. Later,
Schaus’s assignment of M. ostia (and M. parostia comb. n.) to the genus Pamea is strange considering the resemblance of these two species to M. lantona, a species that he described and designated as type species of Menevia, a genus that he also described. In comparing the genitalia of M. ostia with the type species of Pamea, P. albistriga Walker, 1855, the first author found the male genitalia of M. ostia completely unlike those of P. albistriga. The male genitalia of P. albistriga are very simple in comparison with those of M. ostia and Menevia as a whole. The phallus of P. albistriga is simple and truncated apically, and lacking a fused juxta with extended superior processes as in M. ostia and all other Menevia. In P. albistriga the vinculum is much more elongated, valves extremely simple, and the gnathos are atrophied. Genitalia alone offer enough support for the reassignment of P. ostia to Menevia based on the presence of all generic autapomorphies. The external characters of M. ostia are also sufficient for assigning this species to Menevia, namely the postmedial lunule and apical dash, both of which are readily apparent in M. ostia.
Of the four Menevia species-groups, the ostia species-group is the most difficult group to tease apart into different morphologically separable species. Phallic structure is an important character for species identification in Menevia, but each specimen of M. ostia examined (n = 14) had a uniquely shaped dorsal phallic ridge, varying in structure from a singular rounded or rectangular hump, to a crest that follows the length of the phallus (compare Figs
This issue of blurred species boundaries is non-existent in the lantona and lucara species-groups. Both of these species-groups each have one species endemic to the Brazilian Atlantic Forests, and these endemic species each have unique genitalia and display minor but consistent external differences. The high degree of genitalia and external variation in M. ostia, even from a single location, impedes our ability to locate species-specific traits. Molecular evidence may be able to offer more conclusive insights, but even then, material is greatly lacking for the ostia species-group. Until more data is made available, it is more parsimonious to consider M. ostia to be a wide-ranging, phenotypically homogeneous species with variable male genitalia.
Menevia ostia, the very similar M. parostia comb. n., and M. pallida, are apparently distinct on the basis of female morphology and environment, with the latter species being restricted to drier regions of central South America. Only one record of M. ostia exists from a dry region, a specimen from northern Venezuela. The genitalia of this specimen are surprisingly very similar to those of M. pallida. A similar issue exists for the specimens from Espírito Santo. Although these specimens are consistently larger and paler than Costa Rican specimens, some specimens from Costa Rica are in fact, larger and paler than most others from similar localities.
Further complicating our understanding of the distributional boundaries of true M. ostia is a male specimen resembling M. ostia reported from Pook’s Hill, Belize (M. J. C. Barnes pers. comm.; color photo examined). This record is much farther north in Central America than any other records of the ostia group. Only one other species of Menevia is known from Belize: M. menapia. Unfortunately, this specimen is inaccessible to us and it cannot be dissected to determine if it is conspecific with M. ostia, or if it perhaps represents an undescribed species. Regardless of the specific identity of this specimen, the ostia species-group ranges at least as far north as Belize, perhaps making the extreme allopatry of M. menapia less of a biogeographic oddity relative to the genus Menevia as a whole.
Menevia parostia , comb. n.
Pamea parostia Schaus, 1928: 667
Pamea perostia;
Type material
Holotype, ♀: UNKNOWN: Type No. 33592 U.S.N.M./ Pamea parostia type Schaus/
Diagnosis
Menevia parostia can be differentiated from M. ostia by the placement of the postmedial line of the hindwing, which is roughly midway along the length of the hindwing in M. ostia and closer to the wing margin in M. parostia. Furthermore, the sclerotized bands on the venter of the VIII abdominal segment are very thin in M. parostia. Lack of material and variability of this structure, however, belies its diagnostic capability. Additionally, most (93%, n = 15) female specimens of M. ostia are much larger than those of M. parostia. This holotype of M. parostia does not differ remarkably from the single definitive female of M. pallida (see remarks).
Description
Male. Unknown. Female.Head: Straw colored, eyes bordered posteriorly by dark brown collar of scales reaching labial palpi, labial palpi moderately long, reaching beyond frons, segments somewhat well defined ventrally, dorsally with darker scales contrasting with overall lighter coloration. Scape and pedicel weakly tufted. Thorax: As for genus. Straw colored. Legs: As for genus. Tibial spurs relatively thick, long, almost completely scaled except ventrally. Forewing dorsum: Forewing length: 18 mm, n = 1. Subtriangular, rounded, apical quarter of outer margins weakly concave, apex slightly falcate. Ground color pale tan-yellow, moderately speckled by dark petiolate scales. Discal spot faintly marked by gray. Apex marked by black scales near tip of apical dash. Postmedial line brown, mostly straight. Submarginal area pale gray, postmedial lunule originating from near where apical dash meets postmedial line, lunule follows postmedial line from apex to one quarter length of postmedial line where lunule smoothly curves outward toward wing margin becoming somewhat diffuse, forming acute angle with postmedial line. Faint white accessory mark present near tornus. Antemedial line very faint, brown, undulating. Forewing venter: As in forewing dorsum but more heavily speckled, postmedial line bent outwards near tornus, antemedial line absent, discal spot present, small, black. Hindwing dorsum: Rounded, similar coloration and patterning as forewings, postmedial lunule very vague, wavy, not zigzagged, originating near anterior wing margin, following curvature of wing margin, not steeply swept to margin, antemedial area lighter, postmedial line weakly curved, closer to wing margin than midway along wing length. Hindwing venter: Following similar pattern as forewing venter but discal mark absent, marginal area color as surrounding area. Abdomen: As for genus but stouter. Coloration a continuation of thoracic color. Midventral stripe absent. Sternite of VIII as pair of thin sclerotized bands not touching near anterior margin, bowed out slightly mesally. Genitalia: (Fig.
Distribution
Unfortunately the holotype is without locality information, furthermore, Schaus’s original (
Remarks
The presence of such small specimens of M. ostia originally lead us to believe that M. parostia must be just another small example of this species, well within the natural size range and we were prepared to synonymize M. parostia with M. ostia. However, additional examination of the holotype of M. parostia revealed characters of the hindwing maculation and genitalia that were not seen in any examined M. ostia. The paired sternites of VIII in M. parostia are slightly thinner overall in comparison to those of M. ostia, but this character is rather variable in general. A more dramatic difference is found in the arrangement of the hindwing postmedial line between the two species (see Figs
Menevia ostia species-group adults, a recto, b verso. 30 M. ostia ♂, Costa Rica, Guanacaste, Tajo Angeles, 540 m, 10-SRNP-4309 (
Menevia pallida
Menevia pallida Herbin & Mielke, 2014: 147–149; figs ♂ 52–54, ♂ genitalia 55, 56
Type material
Holotype, ♂: BRAZIL: Maranhão: genitalia prep. D. Herbin, ref H. 986 [blue label]/ BRESIL Maranhao [Maranhão], Feira Nova do Maranhão, Retiro, 480 m. 21-25.II.2012, -07°00'31"S, -46°26'41"W, C. Mielke leg. Coll. D. Herbin / DZ 15.734/ Holotype ♂, Menevia pallida Herbin & Mielke det., Antenor 2014 [red label]/ (
Paratype, 1 ♂, BRAZIL: Maranhão: Feira Nova do Maranhão, Retiro, 46°26'41"W, -07°00'31"S, 480 m: 16–17.II.2013, C. Mielke leg., Paratypus Menevia pallida Herbin & C. Mielke det., 2014 [green label], Col. C. Mielke 26.349, St. Laurent diss.: 6-16-15:3 (CGCM).
Additional specimen examined
BRAZIL: Maranhão: 1 ♀, Caixas, Reserva Ecol. Inhamum, 13°12'S, 45°27'W [given coordinates inaccurate], 110 m: 27.II.2006–1.III.2006, lençol e luz mista [sheet and mixed light], F. Limeira-de-Oliveira & J.C. Silva cols, Coleção Embrapa-
Additional questionable specimens examined
(3 ♀ total) BRAZIL: Minas Gerais: 1 ♀, Lassance: 12.XI.1919, Cornel Univ. Expedition Lot 569, Sub 116, Cornell U. Lot 672, Sub 385 Det. W. Schaus [ostia ♀], St. Laurent diss.: 4-20-15:6 (
Diagnosis
Menevia pallida can be differentiated from the somewhat larger, but very similar M. ostia by the pale tan to yellowish brown ground color as opposed to gold or pale yellow. Additionally, the dark speckling is usually heavier due to the presence of more petiolate scales. The hindwings are without bright, obvious, zigzagged postmedial lunules as in M. ostia. In both M. pallida and M. parostia, the placement of the postmedial line on the hindwing is closer to the wing margin than to midway along the length of the wing as it is in M. ostia. The phallic ridge is more smoothly curved and less quadrate terminally than in M. ostia, with the front edge of the phallic ridge evenly sloped rather than squared. The female of M. pallida is smaller than those of M. ostia. We are unable to provide characters to differentiate the unique female of M. parostia from female M. pallida, although the females of M. pallida at our disposal (both true and questionable specimens) are always slightly larger than the unique specimen of M. parostia.
Description
Male.Head: Pale tan, eyes bordered posteriorly by dark brown collar of scales reaching labial palpi, labial palpi small, segments somewhat well defined ventrally, dorsally with darker scales contrasting with overall lighter coloration. Scape and pedicel weakly tufted. Thorax: As for genus. Pale gold-tan. Legs: As for genus. Tibial spurs relatively thick. Forewing dorsum: Forewing length: 15.5–17 mm, avg.: 16.3 mm, n = 2. Triangular, apical half of outer margins weakly concave, apex slightly falcate. Ground color pale tan to yellowish, moderately speckled by dark petiolate scales. Discal spot faintly marked by gray. Apex marked by black scales above apical dash. Black postmedial line mostly straight, sometimes weakly undulating or kinked. Submarginal area pale gray, postmedial lunule originating from near where apical dash meets postmedial line, lunule follows postmedial line from apex to one quarter length of postmedial line where lunule smoothly curves outward toward wing margin becoming somewhat diffuse, forming acute angle with postmedial line. White accessory mark present near tornus. Antemedial line faint, brown, undulating. Forewing venter: As in forewing dorsum but two postmedial lines present, both much fainter than single line on dorsum, one line convex near tornus and slightly undulating, the other straight, following the postmedial line of wing dorsum, antemedial line absent, small black elongated discal spot present. Hindwing dorsum: Rounded, anal angle weakly accentuated, similar coloration and patterning as forewings, postmedial lunule very vague, wavy, not zigzagged, originating near anterior wing margin, following curvature of wing margin, not steeply swept to margin, antemedial area lighter, postmedial line straight, weakly undulating near anal angle. Hindwing venter: Following similar pattern as forewing venter but discal mark absent, marginal area color as surrounding area. Abdomen: As for genus. Coloration a continuation of thoracic color. Midventral stripe absent. Genitalia: (Fig.
Distribution
(Map
Remarks
The recently described M. pallida is not very distinct from the widespread M. ostia or from the unique specimen of M. parostia. The differences between M. pallida and M. ostia, while present, provide only a weak basis on which to consider these taxa as separate species. The original description of M. pallida was based on comparisons with M. lantona, which was considered the most similar species by
Despite the issues with the original description of M. pallida, we still consider it a valid species based primarily on environmental differences, consistently smaller size, paler tan rather than golden coloration, the position of the hindwing postmedial line in females, and the potential difference in female genitalia compared to female M. ostia. The female from Caixas, Maranhão, Brazil, was collected about 440 km northeast of the type locality of M. pallida, and from the same Cerrado habitat (
The single female from Lassance, Minas Gerais, Brazil, which we associate here with M. pallida, albeit questionably, is relatively small and pale compared to M. ostia females from Central America and northern South America, and consequently seems more in line with M. pallida. However, as explained in the remarks to M. ostia, some populations from Atlantic Forest localities are similarly pale. Pending upon the availability of additional specimens of both sexes from Minas Gerais and Espírito Santo, we are unable to conclusively allocate these populations to either species.
We consider two females from Paraguay to be M. pallida due to their localities near the Cerrado, small size, and pale coloration. However, the forewing shape is less rounded than in female M. ostia and M. pallida, making the Paraguayan specimens appear rather distinct (see Fig.
plagiata species-group
The plagiata species-group contains the largest Menevia, the females of some species are among the largest Mimallonidae. Sexual dimorphism is very pronounced in this group. Forewings of females are longer and broader than those of the males, which are conversely very falcate. Unlike in the three previous species-groups, the species belonging to the plagiata group do not have the curved postmedial lunules, but instead have distinct white bands that border the outer margin of the postmedial line, either along the complete length of the line as in the vulgaris subgroup containing: M. vulgaris sp. n., M. franclemonti sp. n., M. vulgaricula sp. n., M. cordillera sp. n., and M. delphinus sp. n.; or are interrupted midway along the line as in the plagiata subgroup, which contains: M. plagiata, M. australis sp. n., and M. alurca. The lack of the lunules immediately distinguishes this species-group from the others. However, the presence of the apical dash is very distinctive and thus the species in this group are easily identifiable as Menevia. Ground color is also darker for this group, most similar to the lucara group, being primarily gray, brown, or some combination. The male genitalia are more robust than in other species-groups. The paired gnathos is flat and oblong, either somewhat triangular or partly ovoid, usually with extreme truncation distally. The phallus is diverse in form, ranging from elongated and smooth to almost triangular due to an exaggerated dorsal projection when viewed laterally. The juxtal arms are generally very flat and wide, not sharp apically.
Key to plagiata species-group
| 1 | Forewing with continuous white band along exterior edge of postmedial line, midventral abdominal stripe absent | 4 [vulgaris subgroup] |
| – | Forewing with interrupted white band along exterior edge of postmedial line, midventral abdominal stripe present | 2 [plagiata subgroup] |
| 2 | Postmedial line usually straight, brown with little red coloration | 3 |
| – | Postmedial line usually bent outwards mesally, coloration dark gray with predominance of red suffusion. Central South America, especially in the Brazilian Cerrado | M. alurca |
| 3 | Wings relatively narrow, usually grayish, white band of forewing postmedial line usually reaching apical tip of wing, especially prominent in females, phallus with elongated, thin, dorsal projection. Vicinity of the state of Rio de Janeiro | M. plagiata |
| – | Wings broader, usually brown, white band of forewing postmedial line usually intercepting apical dash mesally, or dissipating before reaching apex, especially prominent in females, phallus with triangular or rounded dorsal hump. Southeastern Brazil, especially Santa Catarina north to São Paulo | M. australis sp. n. |
| 4 | Large, forewing length males: 22–28 mm, females: 27.5–39 mm, phallus smoothly curving, dorsally smooth or with irregular edge, but never with distinct bulge | 5 |
| – | Small, forewing length males: 17–23 mm, females: 27 mm, phallus with dorsal bulge anteriorly or central protuberance | 6 |
| 5 | Phallus dorsum smooth, evenly edged, terminal sclerotized edge that becomes vesica, diagonal; ventral bump of phallus angled away from phallus terminus. Southeastern Brazil | M. franclemonti sp. n. |
| – | Phallus dorsum with irregular edge, terminal sclerotized edge that becomes vesica nearly vertical, ventral bump of phallus absent or indistinct. Northern South America | M. vulgaris sp. n. |
| 6 | Lobes at base of phallus rounded, not peg-like. Andean or Brazilian Cerrado | 7 |
| – | Lobes at base of phallus peg-like. Amazonian | M. vulgaricula sp. n. |
| 7 | Forewings elongate, very acute apically, forewing postmedial line mostly straight only slightly curved. Andean Cordillera Oriental, Peru to Bolivia | M. cordillera sp. n. |
| – | Forewings stouter, less acute apically, forewing postmedial line slightly undulated. Endemic to the Brazilian Cerrado | M. delphinus sp. n. |
plagiata subgroup
The plagiata subgroup contains M. plagiata, M. australis sp. n., and M. alurca and is diagnosed by the discontinuous white band along the outer margin of the postmedial line and by the presence of a midventral abdominal stripe. The species boundaries in the plagiata subgroup are unclear and thus we consider the three species in this subgroup as part of a species complex hereby-considered the “Menevia plagiata species complex” and may contain additional species not formally recognized herein.
Menevia plagiata
Mimallo plagiata Walker, 1855: 1341
Perophora plagiata;
Menevia plagiata;
Perophora plagiata;
Menevia plagiata;
Menevia plagiata;
Menevia plagiata;
Menevia plagiata;
Menevia plagiata;
Menevia plagiata;
Type material
Holotype, ♂, presumed lost/destroyed. Type locality: Brazil: Rio de Janeiro (see remarks).
Neotype (here designated), ♂: BRAZIL: Rio de Janeiro: BRAZIL: Rio de Janeiro Ste., Teresopolis [Teresópolis], 13–22.iii.1958, H.B.D. Kettlewell, B.M. 1958–273/ NEOTYPE male Mimallo plagiata designated by St Laurent and Dombroskie 2016/ BMNH(E) 1378747/ St. Laurent diss.: 9-2-15:1/ (
Additional specimens examined
(10 ♂, 19 ♀ total) BRAZIL: Espírito Santo: 3 ♀, Santa Teresa: 20.XI.1966, 18.XII.1966, Elias leg. (
Diagnosis
Menevia plagiata is recognizable from all previous species in both sexes by the replacement of the wing margin-swept postmedial lunule with a white band along the length of the postmedial line, which is interrupted midway and resumes near the inner margin. The female of M. plagiata is differentiated from the following two similar species by the presence of a straight or only weakly undulated postmedial line, which, along with the white accessory band, curves toward the apex of the forewing, sometimes sharply, usually to the wingtip, rather than ending at or before reaching the apical dash. Male genitalia are unlike any other species except M. alurca, in that the phallus bears a prominent, elongated, pointed projection from the dorsal surface and is not smooth or ridged as in all other previously diagnosed species. The nearly straight forewing postmedial lines (except near the apex) distinguish both male and female M. plagiata from M. alurca, whereas male genitalia characters and the apical curve of the female forewing postmedial line distinguish M. plagiata from M. australis sp. n.
Description
Male.Head: Gray, eyes large comprising about two-thirds of head area, eyes bordered posteriorly by darker gray collar of scales reaching labial palpi, labial palpi very small, dorsally with darker scales contrasting with overall gray coloration. Scape and pedicel tufted. Thorax: As for genus. Light gray-brown. Legs: As for genus. Tibial spurs small to moderate in length, almost entirely scaled. Forewing dorsum: Forewing length: 22–24.5 mm, avg.: 22.4 mm, n = 7. Triangular, apical half of outer margins concave, apex falcate. Ground color gray-brown with darker gray, brown suffusion especially near interior edge of postmedial line and medial area, reddish coloration near apex along apical interior of postmedial line, overall lightly speckled by dark petiolate scales. Discal spot faintly marked by light gray oblong shape, thin gray mark connecting discal spot to costa. Apex marked by black scales above apical dash, especially near apical tip. Postmedial line straight or weakly undulated, line black, strongly contrasting. Submarginal area light gray with whitish suffusion mesally, postmedial lunule as white band originating from apical dash, white band follows postmedial line from apex to midway along postmedial line, resuming near anal margin. Antemedial line faint, brown, curved outwards. Forewing venter: As in forewing dorsum but grayer rather than brownish, sometimes with pinkish hue, black portion of postmedial line mostly absent except medially where very dark, white outer band of postmedial line as in dorsum, antemedial line absent. Hindwing dorsum: Subtriangular, anal angle weakly accentuated, reddish suffusion near anal angle, similar coloration and patterning as forewings, except postmedial lunule present as zigzagged mark, originating from white outer band along first quarter of postmedial line, postmedial line usually sharply bent toward anterior wing margin, sometimes weakly concave mesally. Hindwing venter: Following similar pattern as forewing venter, but red coloration near anal angle and medial area much darker. Abdomen: As for genus but elongated, nearly sphingiform, reaching beyond anal margin of hindwing. Coloration a continuation of gray thoracic color. Dark, contrasting, midventral stripe present along entire length. Genitalia: (Figs
Distribution
(Map
Natural history
A number of host records exist in the literature for M. plagiatasensu lato, but due to the uncertainty of the identification of this taxon in the past (see remarks below), we cannot be certain that all of the following records pertain to M. plagiatasensu stricto. Perhaps the most reliable record comes from
Remarks
Menevia plagiata is the most problematic taxon in the genus, largely due to the unavailability of the holotype, which is presumed to be lost. The holotype of M. plagiata originated from Fry’s collection, and was collected in Rio de Janeiro.
An additional taxonomic issue was created by the presence of the “holotype” of Perophora ‡superba Jones in the
After visiting the
The taxonomic history of the name plagiata is surely complicated, and unfortunately, without seeing the holotype, we might never be completely sure that our concept of the species coincides with that of
Menevia plagiata species-group [plagiata subgroup] male adults, a recto, b verso. 42 M. plagiata neotype ♂, Brazil, Rio de Janeiro, Teresópolis (
Menevia plagiata species-group [plagiata subgroup] female adults, a recto, b verso. 52 M. plagiata ♀, Brazil, Rio de Janeiro, Petrópolis [photo courtesy CGCM] (
Menevia alurca
Menevia alurcaHerbin & Mielke, 2014: 146–147; figs ♂ 48, 49, ♂ genitalia 50
Menevia ulcara;
Type material
Holotype, ♂: BRAZIL: Maranhão: genitalia prep. D. Herbin, ref H. 1008 [blue label]/ BRESIL Maranhao [Maranhão], Feira Nova do Maranhão, Retiro, 480 m. 4–12.XI.2011, -07°00'31"S, -46°26'41"W, C. Mielke leg. Coll. D. Herbin / Holotype ♂, Menevia alurca Herbin & Mielke det., Antenor 2014 [red label]/ DZ 15.727/ (
Paratypes, 7 ♂ (6 specimens fit our concept of M. alurca, 1 specimen fits our concept of M. delphinus sp. n., see remarks under M. delphinus sp. n.), BRAZIL: Maranhão: 1 ♂, Feira Nova do Maranhão, Retiro, 46°26'41"W, -07°00'31"S, 480 m: 16–17.II.2013, C. Mielke leg., Paratypus Menevia alurca Herbin & C. Mielke det., 2014 [green label], Col. C. Mielke 26.882, St. Laurent diss.: 6-16-15:4 (CGCM) [examined]. Photos of all additional paratypes, each from the same locality and collector as the holotype and single physically examined paratype, were provided by Herbin and Mielke.
Additional specimens examined
(4 ♂, 1 ♀ total) BRAZIL: Distrito Federal: 4 ♂, Estacão Florestal, Cabeça do Veado, 1100 m: 17.X.1971, 19.X.1971, 21.X.1971, 24.X.1971, E.G., I. & E.A. Munroe, St. Laurent diss.: 5-13-15:2 (
Additional transitional and questionable specimens examined
(3 ♂, 1 ♀ total) ARGENTINA: 1 ♀, Formosa: data illegible except for locality,
Diagnosis
Menevia alurca is distinguishable from all previous species in both sexes by the slate gray coloration suffused with deep blood-red, and the postmedial line, which is usually bent outward toward the wing margin at about three-quarters of its length. Additionally, the white band along the exterior of the postmedial line does not curve sharply toward the wing apex as it does in M. plagiata, but instead ends where it meets the apical dash. In this respect, M. alurca is similar to M. australis sp. n.; however, the white band usually juts out sharply just before approaching the apical dash in M. alurca, which is not seen in most other Menevia. The male genitalia however, should immediately distinguish this species from M. australis sp. n.; the phallus bears a prominent, elongated, pointed protuberance from the dorsal surface, not a rounded hump as in M. australis sp.n. The female genitalia differ from those of both M. plagiata and M. australis sp. n. by the small size, the very thin abdominal sclerotizations, and the mesally creased lamella antevaginalis.
Description
Male.Head: Gray, eyes large comprising about two-thirds of head area, eyes bordered posteriorly by darker gray collar of scales reaching labial palpi, labial palpi very small, dorsally with darker scales contrasting with overall gray coloration. Antenna yellowish, scape and pedicel weakly tufted. Thorax: As for genus. Gray. Legs: As for genus. Tibial spurs moderate in length, scaled except for distal quarter. Forewing dorsum: Forewing length: 18–21.5 mm, avg.: 19.7 mm, n = 5. Triangular, apical half of outer margin concave, apex falcate. Ground color dark gray with predominance of deep red-brown or blood-red throughout medial area, overall lightly speckled by dark petiolate scales. Discal spot faintly marked by light gray, oblong shape; thin gray mark connects discal spot to costa. Apex marked by black scales above apical dash. Postmedial line usually bent outwards along three-fourths of its length, rarely nearly straight [especially in transitional population]. Submarginal area light gray, contrasting with much darker medial area, with whitish suffusion mesally forming a faint zigzag, postmedial lunule as distinct white band originating from apical dash, white band follows postmedial line from apex to roughly midway along postmedial line, resuming near anal margin. Antemedial line faint, brown, curved outwards. Forewing venter: As in forewing dorsum but generally much grayer, sometimes with pinkish hue, antemedial line absent, small black discal mark occasionally present. Hindwing dorsum: Subtriangular, anal angle accentuated, reddish coloration usually present near anal angle, bleeding into medial area, similar coloration and patterning overall as forewings, except postmedial lunule present as zigzagged mark, originating from white outer band outlining anterior bend of postmedial line, postmedial line weakly curved toward anterior wing margin, sometimes weakly concave mesally. Hindwing venter: Following similar pattern as forewing venter, but red coloration near anal angle much darker. Abdomen: As for genus, but somewhat stouter. Coloration a continuation of gray thoracic color. Dark, contrasting midventral stripe present. Genitalia: (Fig.
Distribution
(Map
Remarks
Upon reviewing each of the seven paratypes of M. alurca, it became apparent that one paratype in the collection of Herbin (Bc-Her4848) does not belong to this species and was incorrectly included in the paratype series. The specimen in question has a continuous white band along the forewing postmedial line and lacks a midventral stripe on the abdomen, characters that allow us to identify this specimen as M. delphinus sp. n., another Cerrado species, described below.
External characteristics and male genitalia, namely, the phallus, of M. alurca is extremely similar to that of M. plagiata, thus these species are not readily differentiable based on male genitalia alone, as are most Menevia species. Furthermore, three specimens from two nearby localities in central Minas Gerais that we questionably determined as M. alurca, are somewhat intermediate in habitus between M. alurca from farther north and west (central Cerrado), and M. plagiata from farther east and southeast (Brazilian Atlantic Forest). These specimens display the predominance of darker coloration and the arrangement of markings near the forewing apices that we attribute to M. alurca. However, the forewing postmedial line is only weakly curved outwards, not bent so dramatically as in the types of M. alurca. These specimens are also quite large, much more in line with those of M. plagiata. The population from Minas Gerais is transitional in appearance between M. alurca and M. plagiata, and it is worth noting that the Cerrado and Atlantic Forest biomes converge there (
A female that we questionably assign to this species, from Formosa, Argentina, is extremely small and has a nearly straight forewing postmedial line. Although the size and postmedial line are not comparable to the other female M. alurca, the markings at the forewing apex, the predominance of red coloration, and genitalia (namely the mesally creased lamella antevaginalis), are all highly suggestive of M. alurca. Without more material from this locality, particularly males, it is impossible to reach conclusive determination about its identity. We decided to include this specimen in our material examined due to its collecting locality, which represents the only Menevia record from Argentina.
Menevia australis sp. n.
Type material
Holotype, ♂: BRAZIL: Santa Catarina: Jaragua [Jaraguá do Sul], Santa Catharina, Brazil, 5 Sept 1934, Fritz Hoffmann/ PARATYPE Menevia elegans J. G. Franclemont/ St. Laurent diss.: 6-19-15:4/ Franclemont’s label refers to a MS name./ HOLOTYPE male Menevia australis St Laurent and Dombroskie, 2016 [handwritten red label]/ (
Paratypes, 19 ♂, 9 ♀: BRAZIL: Santa Catarina: 3 ♂, São Bento do Sul, Rio Vermelho, 850 m: 16.IX.1985, Mielke, Rank & Casagrande leg., DZ 32.713–32.715 (
Diagnosis
Menevia australis is similar to M. plagiata in both sexes, but can be distinguished in both sexes by slightly broader wings, deeper brown coloration, and by the white band along the postmedial line, which terminates at the apex somewhat mesally along the length of the apical dash. The white band is either nearly perpendicular to the apical dash or it forms a roughly 45 degree angle with it. The white band may be somewhat curved toward the apex as in M. plagiata, but does not reach the apical tip of the wing. Male genitalia are easily recognized by the rounded hump or short triangular protuberance on the dorsal surface of the phallus, easily distinguishing the male genitalia from those of M. plagiata and M. alurca, which both have very elongated dorsal protuberances on the phallus. Additionally, the uncus is narrower and more triangular in M. australis. The VIII tergite in the female genitalia of M. australis forms a rounded arc and is not triangular as in M. plagiata and M. alurca.
Description
Male.Head: Gray-brown or light brown, eyes large comprising about two-thirds of head area, eyes bordered posteriorly by darker brown collar of scales reaching labial palpi, labial palpi very small, dorsally with darker scales contrasting with overall gray coloration. Scape and pedicel tufted. Thorax: As for genus. Light gray-brown to light brown. Legs: As for genus. Tibial spurs small to moderate in length, almost entirely scaled. Forewing dorsum: Forewing length: 21.5–23 mm, avg.: 22.4 mm, n = 7. Triangular, apical half of outer margin concave, apex falcate. Ground color gray-brown with caramel brown or almost slate gray suffusion throughout medial area, reddish coloration near apex along apical interior of postmedial line, overall lightly speckled by dark petiolate scales. Discal spot faintly marked by light gray oblong shape, thin gray mark connecting discal spot to costa. Apex marked by black scales above apical dash, especially near apical tip. Postmedial line straight or weakly undulated, line black, strongly contrasting. Submarginal area light gray with whitish suffusion mesally forming faint or conspicuous zigzag, postmedial lunule as white band originating mesally from apical dash, white band follows postmedial line from apex to midway along postmedial line becoming zigzagged diffusion, white band resumes near anal margin. Antemedial line faint, brown, curved outwards. Forewing venter: As in forewing dorsum but grayer rather than brownish, antemedial line absent. Hindwing dorsum: Subtriangular, anal angle weakly accentuated, reddish coloration usually present near anal angle, similar coloration and patterning as forewings, except postmedial lunule present as zigzagged mark similar to zigzagged diffusion on forewing, mark originating from white outer band along first quarter of postmedial line, postmedial line sharply bent toward anterior wing margin, sometimes weakly concave mesally. Hindwing venter: Following similar pattern as forewing venter, but red coloration near anal angle much darker, almost brown. Abdomen: As for genus but somewhat elongated, nearly sphingiform. Coloration a continuation of gray thoracic color. Dark, contrasting midventral stripe present. Genitalia: (Fig.
Distribution
(Map
Etymology
Menevia australis is named for its southerly distribution, which among Menevia, is only shared with M. magna. Additionally, M. australis seems to represent the southeast most extension of the M. plagiata species complex, replacing M. plagiata farther southeast.
Remarks
Menevia australis is the southeasternmost species of the M. plagiata complex and is quite difficult to separate from true M. plagiata without a genitalia dissection or geographic information. However, upon thorough examination of the male and female genitalia, external diagnostic characters became readily apparent and have been presented above in the diagnosis. Additionally, the allopatric distribution of these species suggests that they are two separate species, albeit very closely related. The allopatric distribution patterns of M. australis and M. plagiata are not unique, similar allopatry was shown in two closely related Saturniidae by
Both M. australis and M. franclemonti sp. n. described below, were originally recognized as distinct by J. G. Franclemont, and given manuscript names, but never formally described. The holotype and some paratypes of M. australis (all from Jaraguá do Sul, Santa Catarina, Brazil) bear labels reading “PARATYPE,” “HOLOTYPE,” or “ALLOTYPE” with Franclemont’s manuscript name Menevia ‡elegans. In addition to our holotype and paratype labels, we have placed labels on these specimens stating that Franclemont’s labels represent a manuscript name.
vulgaris subgroup
The vulgaris subgroup contains M. vulgaris sp. n., M. franclemonti sp. n., M. vulgaricula sp. n., M. cordillera sp. n., and M. delphinus sp. n. and is diagnosed by the continuous white band along the outer margin of the postmedial line and by the lack of a midventral abdominal stripe. The species boundaries in the vulgaris subgroup are clearer than in the previous subgroup.
Menevia vulgaris sp. n.
Type material
Holotype, ♂: FRENCH GUIANA: GUYANE, St. Laurent-du-Maroni, Piste de Paul Isnard, N05°22.562', W53°57.678', 8.iii.2011, 52 m., P. Sammut/ MENEVIA plagiataplagiata (Walker, 1855)/ St. Laurent diss.: 9-2-15:2/ HOLOTYPE male Menevia vulgaris St Laurent and Dombroskie, 2016 [handwritten red label]/ (
Paratypes, 30 ♂, 10 ♀: BRAZIL: Amazonas: 8 ♂, Reserva Ducke, km. 26 Manaus-Itacoatiara Highway: 15.IV.1972, 16.IV.1972, 19.IV.1972, 15.V.1972, 16.V.1972, E.G., I. & E.A. Munroe, St. Laurent diss.: 4-25-15:8, 8-3-15:1 (
Diagnosis
Menevia vulgaris is recognizable from all previous species by the replacement of the wing margin swept postmedial lunule with a continuous white band along the entire length of the postmedial line. This species is quite large for the genus, with highly elongated acutely triangular forewings and triangular hind wings. Sexual dimorphism is well developed, with females having broader, less triangular, but still highly elongated forewings. Male genitalia are easily recognized by the somewhat cylindrical shape of the phallus with an irregularly edged dorsum lacking an extensive dorsal ridge, but usually with a weak anteriorly situated bulge. The juxtal processes are wide and flattened, but only weakly sclerotized; the proximal lobes of the phallus are also very broad, not elongated or fingerlike as in most other species. The combination of these genitalia characters also distinguish M. vulgaris from the following four new species described below. Size should also be sufficient in separating M. vulgaris from all sympatric species, as M. vulgaris is always the largest species within its range. Menevia vulgaris is most similar in appearance and size to M. franclemonti sp. n. described below, but is easily differentiated from this species by the more sharply angled hindwing postmedial line and by the more uneven dorsum of the phallus. Geography should also be sufficient for separating M. vulgaris from M. franclemonti sp. n., as the latter species is restricted to southeastern Brazil, outside the largely Amazonian distribution of M. vulgaris.
Description
Male.Head: Gray, eyes large comprising about two-thirds of head area, eyes bordered posteriorly by darker gray collar of scales reaching labial palpi, labial palpi very small, segments weakly defined ventrally, dorsally with darker scales contrasting with overall gray coloration. Scape and pedicel tufted. Thorax: As for genus. Light gray. Legs: As for genus. Tibial spurs very small, short, almost entirely scaled. Forewing dorsum: Forewing length: 22–28 mm, avg.: 25.3 mm, n = 24. Elongated, acutely triangular, apical half of outer margin concave, apex falcate. Ground color gray with darker gray, brown, or reddish brown suffusion throughout medial area, especially near interior edge of postmedial line, pinkish red to blood red coloration near apex along apical interior of postmedial line, overall lightly speckled by dark petiolate scales. Discal spot faintly marked by light gray oblong shape, thin gray mark connecting discal spot to costa. Apex marked by black scales above extended apical dash. Postmedial line weakly curved to follow outline of wing margin, line black, strongly contrasting. Submarginal area light gray with whitish suffusion mesally, postmedial lunule as distinct white band originating from apical dash, white band follows postmedial line from apex to posterior wing margin. Antemedial line faint, brown, curved outwards. Forewing venter: As in forewing dorsum but antemedial area lighter gray, more contrasting, sometimes with blood-red suffusion, antemedial line absent, small black discal spot occasionally present. Hindwing dorsum: Triangular, anal angle weakly accentuated, reddish coloration near anal angle, similar coloration and patterning as forewings, antemedial line absent, postmedial line sharply bent toward anterior wing margin, concave mesally. Hindwing venter: Following similar pattern as forewing venter, but red coloration near anal angle much darker, discal mark sometimes present. Abdomen: As for genus, but somewhat elongated, nearly sphingiform. Coloration a continuation of gray thoracic color. Midventral stripe absent. Genitalia: (Fig.
Distribution
(Map
Etymology
Menevia vulgaris (=vulgaris Latin, meaning commonplace) is named for its wide distribution and apparent commonness.
Remarks
This species is usually the most frequently represented Menevia in collections and apparently quite common throughout its broad distribution. This new species is usually misidentified as M. plagiata and has long been considered conspecific with the southeastern Brazilian species redescribed above. The taxonomic issues surrounding the name plagiata have been explained in the remarks of that species, and the designation of the neotype of M. plagiata resolves any previous identification problems and allows recognition of these two very distinct taxa.
Currently, the name M. vulgaris can now only be associated with the large species present in the Amazonian and Cerrado regions. Diagnostic characters given before, particularly size and male genitalia, are adequate for identifying this species. Other names, including plagiata and the various new species described below, are associated with either allopatric or much smaller species.
As with other wide-ranging Menevia species, such as M. lantona and M. lucara, there is a degree of geographic variation, although this variation is less obvious than in these two species. As in other species-groups, M. vulgaris from Colombia have more robust male genitalia, except that the juxtal processes are thinner, the valve teeth smaller, and the gnathos elongations shorter. Externally, however, the examined males do not differ from other populations except for their overall slightly larger size. Similar shortened gnathos elongations were found in the males from Ecuador. These males and a single examined female from Ecuador were all smaller than males and females from other populations. However, the lack of differences of the phallus in Ecuadorian males compared to those from other locations suggests that the smaller size and some noted differences found in the single female dissection, namely the shape of the highly variable ventral sclerotized bands on VIII, are not grounds to consider the Ecuadorian population a distinct taxon. Another variation worth noting is in the single Peruvian female specimen, which has the reddish maculation on all wings replaced by deeper brownish-red.
Menevia plagiata species-group [vulgaris subgroup] male adults, a recto, b verso. 58 M. vulgaris holotype ♂, French Guiana, St. Laurent du Maroni, Piste de Paul Isnard, 52 m (
Menevia plagiata species-group [vulgaris subgroup] female adults, a recto, b verso. 67 M. vulgaris paratype ♀, Guyana, Omai (
Menevia male genitalia, a ventral, b lateral, c phallus [except Figures 80, 81]. 72 M. lantona, French Guiana, Pied Saut, Oyapok River [St. Laurent diss.: 3-7-15:2] (CMNH) 73 M. rosea holotype, Ecuador, Napo, Simón Bolívar, 1200 m [St. Laurent diss.: 3-7-15:9, 73b inverted horizontally] (CMNH) 74 M. torvamessoria holotype, Peru, Puno, La Unión, 2000 ft [St. Laurent diss.: 6-29-15:4, 74b inverted horizontally] (
Menevia male genitalia, a ventral, b lateral, c phallus. 82 M. pallida paratype, Brazil, Maranhão, Feira Nova do Maranhão, 480 m [St. Laurent diss.: 6-16-15:3] (CGCM) 83 M. plagiata neotype, Brazil, Rio de Janeiro, Teresópolis [St. Laurent diss.: 9-2-15:1] (
Menevia male genitalia, a ventral, b lateral, c phallus. 87 M. vulgaris holotype, French Guiana, St. Laurent du Maroni, Piste de Paul Isnard, 52 m [St. Laurent diss.: 9-2-15:2] (
Menevia female genitalia, a ventral, b VIII tergite detail. 92 M. lantona, French Guiana, St. Jean du Maroni, [St. Laurent diss.: 7-8-15:1] (
Menevia female genitalia, a ventral, b VIII tergite detail. 98 M. plagiata, Brazil, Rio de Janeiro [St. Laurent diss.: 7-28-15:1] (
Menevia female genitalia, a ventral, b VIII tergite detail. 101 M. vulgaris paratype, French Guiana, St. Laurent du Maroni [St. Laurent diss.: 4-25-15:13] (
Menevia franclemonti sp. n.
Type material
Holotype, ♂: BRAZIL: Santa Catarina: Jaragua [Jaraguá do Sul], Santa Catharina, Brazil, 10 Nov 1934, Fritz Hoffmann/ PARATYPE Menevia falco J. G. Franclemont/ St. Laurent diss.: 4-25-15:5/ Franclemont’s label refers to a MS name./ HOLOTYPE male Menevia franclemonti St Laurent and Dombroskie, 2016 [handwritten red label]/ (
Paratypes, 10 ♂, 1 ♀: BRAZIL: Rio de Janeiro: 1 ♂, Petrópolis, Independência, 900 m: 15.XI.1939, Gagarin leg., ex. col. Gagarin, DZ 32.717 (
Diagnosis
Very similar to M. vulgaris, but distinguished by the usually warmer brown hue rather than gray, more sharply falcate forewings, and a more smoothly curving postmedial line on the hindwing, which does not sharply bend toward the anterior wing margin. The male genitalia are also unique in that the phallus is thin and smooth, not irregularly shaped dorsally, with the sclerotized portion of the distal end of the phallus diagonal rather than vertical as in M. vulgaris. Furthermore, the base of the sclerotized terminus of the phallus, on the venter, has a prominent bump, angled away from the distal end of the phallus. Menevia franclemonti replaces M. vulgaris in the Brazilian Atlantic Forest.
Description
Male.Head: Gray, eyes large comprising about two-thirds of head area, eyes bordered posteriorly by light brownish gray collar of scales reaching labial palpi, labial palpi very small, dorsally with darker scales contrasting with overall gray coloration. Scape and pedicel tufted. Thorax: As for genus. Light gray to brown. Legs: As for genus. Tibial spurs small, short, almost entirely scaled, except distal tip. Forewing dorsum: Forewing length: 25–27 mm, avg.: 26.1 mm, n = 6. Elongated, acutely triangular, apical half of outer margin highly concave, apex very falcate. Ground color gray or warm brown with darker gray, brown, or reddish brown suffusion, especially near interior edge of postmedial line and medial area; brownish red to blood red coloration near apex along apical interior of postmedial line, overall lightly speckled by dark petiolate scales. Discal spot faintly marked by light gray oblong shape, thin gray mark connecting discal spot to costa. Apex marked by black scales above extended apical dash. Postmedial line smooth, mostly straight, line black, strongly contrasting. Submarginal area light gray with small grayish-white suffusion mesally, postmedial lunule as white band originating from apical dash, white band follows postmedial line from apex to anal wing margin. Antemedial line faint, brown, curved outwards. Forewing venter: As in forewing dorsum but antemedial area lighter gray, more contrasting, blood-red suffusion more pronounced, antemedial line absent, small black discal mark occasionally present. Hindwing dorsum: Triangular, anal angle usually sharply accentuated, reddish coloration that bleeds into medial area originates near anal angle, overall similar coloration and patterning as forewings, antemedial line absent, postmedial line weakly curved toward anterior wing margin, not concave mesally. Hindwing venter: Following similar pattern as forewing venter, but red coloration near anal angle much darker, discal mark sometimes present. Abdomen: As for genus, but somewhat elongated, nearly sphingiform. Coloration a continuation of gray thoracic color. Midventral stripe absent. Genitalia: (Fig.
Distribution
(Map
Etymology
Menevia franclemonti is named after the lepidopterist J. G. Franclemont, who originally recognized the uniqueness of this species. He also wrote an important fascicle on the Mimallonidae of North America north of Mexico (
Remarks
This new species is the Brazilian Atlantic Forest component of the vulgaris subgroup of the plagiata species-group in much the same way that M. mielkei and M. magna are the Atlantic Forest components of the lucara and lantona species-groups respectively. Although M. franclemonti is not remarkably distinct from M. vulgaris, it is readily differentiated by the external and genitalia diagnostic characters presented above. Additionally, this species seems to be allopatric to all other species in this subgroup. Despite the differences, M. franclemonti, together with the similar M. vulgaris, have both been misidentified as M. plagiata for the reasons explained in the remarks of M. plagiata and M. vulgaris. The allopatry of M. vulgaris and M. franclemonti certainly affords that M. plagiatasensu stricto could not be applied to the Amazonian M. vulgaris, but it is certainly plausible that
As in the case of M. australis, J. G. Franclemont recognized this species as distinct and labeled specimens as holotype and paratypes (all from Jaraguá do Sul, Santa Catarina, Brazil) under his manuscript name Menevia ‡falco. Rather than naming this species M. ‡falco as he originally intended, we decided to honor J. G. Franclemont by naming this species after him for his work on Menevia and Mimallonidae in general.
Menevia vulgaricula sp. n.
Type material
Holotype, ♂: BRAZIL: Amazonas: Hyutanahan [Huitanaã], Rio Purus, Brazil, S.M. Klages/ Jan. 1922/ Carn. Mus. Acc. 6963/ St. Laurent diss.: 6-19-15:1/ HOLOTYPE male Menevia vulgaricula St Laurent and Dombroskie, 2016 [handwritten red label]/ (CMNH). Type locality: Brazil: Amazonas: Huitanaã.
Paratypes, 2 ♂, 1 ♀: BRAZIL: Amazonas: 1 ♂, Hyutanahan [Huitanaã], Rio Purus: II.1922, S.M. Klages, Carn. Mus. Acc. 6963, St. Laurent diss.: 4-25-15:14 (CMNH). 1 ♂, Nova Olinda, Rio Purus: V.1922, S.M. Klages, St. Laurent diss.: Carn. Mus. Acc. 6962, St. Laurent diss.: 6-19-15:2 (CMNH). 1 ♀, Rio Madeira: Juillet-Août (Fassl), Dognin Collection,
Additional questionable specimens examined
(3 ♀ total) [not included in type series] BRAZIL: Pará: 1 ♀, Likely Belém: A.M. Moss, Rothschild Bequest 1939–1, St. Laurent 7-21-15:2, BMNH(E) 1378755 (
Diagnosis
Menevia vulgaricula is similar to the previous two species, but much smaller, both males and females are notably smaller than the respective sexes of M. vulgaris. The female of M. vulgaricula is easily differentiated from females of M. vulgaris by the width of the submarginal area, which is broader and decreases in width toward the apex much less gradually than the rapidly narrowing submarginal area of M. vulgaris. The genitalia of both sexes can be used to differentiate M. vulgaricula from similar species (except from the unexamined female of M. franclemonti). In males, the phallus has a very prominent setae covered dorso-anterior bulge reminiscent of a dorsal phallic ridge, the tegumen, vinculum, and acutely triangular uncus are all very narrow and elongated, and the paired processes of the gnathos converge and bend upwards. Additionally, the divergent lobes at the proximal end of the phallus are not broad, but thin and peg-like. In the female, the lamella antevaginalis is very thin, unlike the usually broad, quadrate lamella of M. vulgaris.
Description
Male.Head: Gray, eyes large comprising about two-thirds of head area, eyes bordered posteriorly by brownish gray collar of scales reaching labial palpi, labial palpi very small, segments weakly defined ventrally, dorsally with darker scales contrasting with overall gray coloration. Scape and pedicel tufted. Thorax: As for genus. Light gray. Legs: As for genus. Tibial spurs moderate length, thin, somewhat hooked distally. Forewing dorsum: Forewing length: 19–22 mm, avg.: 21 mm, n = 3. Acutely triangular, apical half of outer margin concave, apex falcate. Ground color gray with brown or reddish brown suffusion, especially near interior edge of postmedial line and medial area, reddish coloration near apex along apical interior of postmedial line, overall lightly speckled by dark petiolate scales. Discal spot faintly marked by light gray oblong shape, thin gray mark connecting discal spot to costa. Apex marked by black scales above extended apical dash. Postmedial line mostly straight except when approaching apex, line black, strongly contrasting. Submarginal area light gray with whitish suffusion mesally, sometimes appearing as a faint zigzag, postmedial lunule as white band originating from apical dash, white band follows postmedial line from apex to anal wing margin. Antemedial line faint, brown, curved outwards. Forewing venter: As in forewing dorsum but antemedial area lighter gray, more contrasting, blood red suffusion more expansive, antemedial line absent, small black discal spot occasionally present. Hindwing dorsum: Triangular, anal angle weakly accentuated, reddish coloration near anal angle, similar coloration and patterning as forewings, antemedial line absent, postmedial line moderately bent toward anterior wing margin, weakly concave mesally. Hindwing venter: Following similar pattern as forewing venter, but red coloration near anal angle much darker, discal mark absent or very faint. Abdomen: As for genus but somewhat elongated, nearly sphingiform. Coloration a continuation of gray thoracic color. Midventral stripe absent. Genitalia: (Fig.
Distribution
(Map
Etymology
Menevia vulgaricula is named for its appearance as a diminutive M. vulgaris.
Remarks
Menevia vulgaricula is an interesting species due to its remarkable external resemblance to M. vulgaris, with which M. vulgaricula is sympatric. Despite notable similarity in external characters, M. vulgaricula is much smaller and displays distinct genitalia characters, in both the male and the female.
Both M. vulgaris and M. vulgaricula were collected by S. M. Klages during the same period at both Huitanaã and Nova Olinda, Amazonas, Brazil (CMNH). Therefore, not only are these two species sympatric, but are also apparently both active at the same time of year. Size and distinct differences in genitalia likely afford some form of a prezygotic barrier. The last two new species described below are very similar to M. vulgaricula, in that each species is very small relative to the larger M. vulgaris and M. franclemonti, and both display prominent anterior phallic bulges. These species however are widely allopatric. Therefore, the modes of isolation between sympatric M. vulgaris and M. vulgaricula certainly warrants future investigation.
Additional females from Pará, Brazil and French Guiana were examined, but cannot be included in the type series because we lack the more easily identifiable males from these localities. Also, these specimens display deeper red coloration and wavier forewing postmedial lines, characters not seen in all other examined M. vulgaricula. Although the small size of these females and the genitalia of the Pará specimen suggest that they are probably M. vulgaricula, Pará and French Guiana are distant from both Rio Madeira and Rio Purus, the two relatively nearby localities known to support M. vulgaricula.
Menevia cordillera sp. n.
Type material
Holotype, ♂: PERU: 1584, San Gaban [San Gabán], Peru, 2500 ft, March–April 1913/ Joicey Coll. Brit. Mus. 1925–157/ St. Laurent diss.: 6-29-15:8/ BMNH(E) 1378757/ HOLOTYPE male Menevia cordillera St Laurent and Dombroskie, 2016 [handwritten red label]/ (
Paratype, 1 ♂: BOLIVIA: Rio Songo [Río Zongo], 750 m: Coll. Fassl, Dognin Collection,
Diagnosis
This new species, like M. vulgaricula, is quite small in comparison with the widespread M. vulgaris and the southeast Brazilian M. franclemonti. Due to the small size of M. cordillera, it may be confused with the allopatric M. vulgaricula but can easily be differentiated by the deeper reddish brown coloration, more sharply acute apices of the more elongated forewings, straighter hindwing margins, and by the male genitalia. The phallus of M. cordillera is somewhat reminiscent of that of M. vulgaricula, but with a more triangularly shaped anterior dorsal bulge and rounded, not peg-like, lobes at the base of the phallus. Overall, the phallus of M. cordillera is broader than that of M. vulgaricula. No other Menevia species are known from the Cordillera Oriental, besides the clearly distinct M. torvamessoria.
Description
Male.Head: Light brown-gray, eyes large comprising about two-thirds of head area, eyes bordered posteriorly by dark brown collar of scales reaching labial palpi, labial palpi large, robust for genus, dorsally with darker scales contrasting with overall gray coloration. Scape and pedicel weakly tufted. Thorax: As for genus. Light tan. Legs: As for genus. Tibial spurs short, stout. Forewing dorsum: Forewing length: 22–23 mm, avg.: 22.5 mm, n = 2. Very acutely triangular, apical half of outer margin deeply concave, apex very falcate. Ground color gray with deep reddish brown suffusion throughout medial area, brighter reddish coloration near apex along apical interior of postmedial line, overall lightly speckled by dark petiolate scales. Discal spot faintly marked by light gray oblong shape, gray mark connecting discal spot to costa. Apex marked by black scales above extended apical dash. Black postmedial line mostly straight except when approaching apex where sharply curved, strongly contrasting. Submarginal area light gray with whitish suffusion mesally, sometimes appearing as faint zigzag, submarginal area with distinct white band originating from apical dash, white band follows postmedial line from apex to anal wing margin. Antemedial line very faint, brown, curved outwards. Forewing venter: As in forewing dorsum but antemedial area lighter gray, more contrasting, antemedial line absent, small black discal mark present. Hindwing dorsum: Triangular, outer margin very straight, anal angle weakly accentuated, reddish coloration near anal angle, bleeding into medial area, similar coloration and patterning as forewings, antemedial line absent, postmedial line sharply bent toward anterior wing margin, weakly concave mesally. Hindwing venter: Following similar pattern as forewing venter, but red coloration near anal angle much darker, discal mark absent. Abdomen: As for genus, but elongated, nearly sphingiform. Midventral stripe absent. Genitalia: (Fig.
Distribution
(Map
Etymology
This new species is named for the Andean Cordillera Oriental, to which this species is endemic.
Remarks
Menevia cordillera is very closely related to both M. vulgaris and M. vulgaricula based on general external characters and the genitalia morphology. These three species may represent taxa of a species complex that spans throughout northern South America.
An additional specimen from the Yungas of Bolivia, in the collection of Daniel Herbin (Bc-Her2532) as seen in the BOLD database, almost certainly belongs to this new species. However, we were unable to examine this specimen and thus it cannot be included in the type series, but we report it here as it provides additional distributional data.
Menevia delphinus sp. n.
Type material
Holotype, ♂: BRAZIL: Distrito Federal: Coleção EMBRAPA-
Paratypes, 6 ♂, 2 ♀: BRAZIL: Distrito Federal: 3 ♂, 2 ♀, Planaltina, 15°35'S, 47°42'W, 1000 m: 11.XI.1977, 15.XI.1977, V.O. Becker col., Col. Becker No. 22269, 22304, St. Laurent diss.: 4-25-15:16,
Additional specimen examined from photo
[not included in type series] BRAZIL: Maranhão: 1 ♂, Feira Nova do Maranhão, Retiro, 46°26'41"W, -07°00'31"S, 480 m: C. Mielke leg., Paratype, Menevia alurca Herbin & Mielke, 2014 (Collection of Daniel Herbin, France).
Diagnosis
Menevia delphinus, like the previous two species, is quite small in comparison with the widespread M. vulgaris and the southeast Brazilian M. franclemonti, but males and females can easily be differentiated from all others belonging to the plagiata species-group by the relatively stout forewings and by the genitalia. The phallus of M. delphinus is most similar to that of M. cordillera, but with the dorsal bulge being situated more mesally along the length of the phallus and usually much more pronounced as a singular, blunt, protuberance, not a pointed or triangular bulge as in other species. Additionally, the lobes at the base of the phallus in M. delphinus are very elongated and tubular, almost fingerlike, not rounded or peg-like as in the previous similar species. In females, the VIII tergite is very robust, forming a distinct triangle, and is not rounded and arc-like as in some similar species. Menevia delphinus may also be confused with M. alurca due to the similar size and sympatry; however, M. delphinus can be straightforwardly recognized by the lack of a midventral abdominal stripe and a continuous white band along the outer edge of the postmedial line, which is discontinuous midway along the postmedial line in M. alurca. Additionally, the genitalia easily differentiate M. delphinus from M. alurca. The dorsal protuberance of the phallus of M. delphinus is much smaller and blunter in comparison with the extremely elongate, curved, and sharply pointed dorsal phallic protuberance of M. alurca. The only other sympatric species besides M. alurca is the much larger M. vulgaris.
Description
Male.Head: Light brown or gray, eyes large comprising about two-thirds of head area, eyes bordered posteriorly by brownish collar of scales reaching labial palpi, labial palpi very small, short, covered in dark scales. Scape and pedicel weakly tufted. Thorax: As for genus. Light gray. Legs: As for genus. Tibial spurs moderate length, thin, scaled except for distal tip. Forewing dorsum: Forewing length: 17–23 mm, avg.: 20.5 mm, n = 6. Triangular, not overly elongated, apical half of outer margin concave, apex falcate. Ground color gray with reddish brown suffusion, especially near interior edge of postmedial line, reddish coloration near apex along apical interior of postmedial line, overall moderately speckled by dark petiolate scales. Discal spot very faintly marked by light gray oblong shape. Apex marked by black scales above apical dash. Black postmedial line mostly straight except when very near apex, strongly contrasting. Submarginal area light gray with whitish suffusion mesally, postmedial lunule as white band originating from apical dash, white band follows postmedial line from apex to posterior wing margin. Antemedial line brown, almost nonexistent. Forewing venter: As in forewing dorsum but antemedial area lighter gray, blood red suffusion present, especially along interior edge of postmedial line and near apex, antemedial line absent, small black discal mark occasionally present. Hindwing dorsum: Triangular, anal angle weakly accentuated with reddish coloration, similar coloration and patterning as forewings, antemedial line absent, postmedial line weakly bent toward anterior wing margin, weakly concave or straight mesally. Hindwing venter: Following similar pattern as forewing venter, but red coloration near anal angle much darker, discal mark absent or very faint. Abdomen: As for genus but somewhat elongated, nearly sphingiform. Coloration a continuation of gray thoracic color. Midventral stripe absent. Genitalia: (Fig.
Distribution
. This new species is a resident of the Brazilian Cerrado in the states of Goiás and Minas Gerais, as well as in Distrito Federal. An additional record from Maranhão will be discussed below in the remarks.
Etymology
This new species is named for the phallus, which, due to the dorsal protuberance, bears the likeness of a dolphin (=delphinus Latin) when viewed laterally.
Natural history
Dr. A. Camargo (
Remarks
Menevia delphinus represents another species similar to the large, widespread M. vulgaris and its Brazilian Atlantic Forest counterpart, M. franclemonti. Menevia delphinus, like three other similar species described as new in the present work, is much smaller and with much more complicated phallic structures than M. vulgaris and M. franclemonti.
Due to the lack of data regarding the distribution of this species, other than it clearly being found in the Brazilian Cerrado, we consider it is necessary to report another state record for this species, despite our inability to gain access to the specimen in question. The recently described, and very distinct, M. alurca, was described from eight males (
The single reared specimen of M. delphinus was much smaller than other M. delphinus specimens and its genitalia differed in the shape of the dorsal protuberance of the phallus, which was flattened rather than distinctly raised. However, other genital characteristics, such as the highly elongated lobes at the base of the phallus, were consistent with M. delphinus. Given the consistency in other characters and the close proximity of this specimen’s locale to the type locality of M. delphinus, we include this specimen in the type series and attribute the different phallic structure to the overall small size.
Mimallo saturata
Type material
Holotype, ♀, presumed lost/destroyed. Type locality: Brazil: Rio de Janeiro (see remarks of Menevia plagiata).
The unknown taxon Mimallo saturata was briefly discussed above in the remarks relating to Menevia plagiata because some specimens of Menevia had previously been attributed to Mimallo saturata (
Additional discussion
Menevia is a wide-ranging genus displaying distinct patterns of speciation. For example, both the lantona and lucara species-groups are widely sympatric, with exceptions only in Central America. In both species-groups we discovered taxa distinct from the nominotypical species on the peripheries of those species’ distributions, in southeastern Brazil, Central America, and the Andean Cordillera Oriental. The plagiata species-group also showed similar patterns of speciation in the Andean Cordillera Oriental and southeastern Brazil. Additionally, the plagiata and ostia species-groups both have unique Brazilian Cerrado species. The broad overlap in the distributions of the four species-groups suggests that they may have originated from the same geographic region, with various degrees of parallel speciation.
The distinct dorsal phallic ridge or projection of the male genitalia is an important character used to differentiate species in this genus, and interestingly, all species from southeastern Brazil, except M. franclemonti, display this character, whereas all Amazonian and Central American species, except M. ostia, do not. Additionally, the midventral abdominal stripe is present only in central and southeastern Brazilian species, namely in M. plagiata, M. alurca, M. australis, and M. magna. This trait is not present in Amazonian or Central American species. Most species displaying the midventral abdominal stripe belong to a closely-knit section of the M. plagiata subgroup of the plagiata species-group, therefore its presence in M. magna of the lantona species-group implies that this trait may be plesiomorphic along with dorsal ridges/projections of the phallus which this species also displays. Furthermore,
A clearer understanding of the evolutionary origins of Menevia, and Mimallonoidea as whole, would be particularly interesting because recent higher-level phylogenetic studies of Lepidoptera continue to demonstrate not only the uniqueness of Mimallonidae as the only family in Mimallonoidea, but also its key phylogenetic role as a potential sister lineage to the Macrolepidoptera (
Acknowledgements
We thank the following individuals for access to collections and facilitation of specimen loans: J. Barbut (
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