Research Article |
Ryan A. St Laurent‡, Daniel Herbin§, Carlos G. C. Mielke|
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Corresponding author: Ryan A. St Laurent ( rstlaurent@flmnh.ufl.edu ) Academic editor: Donald Lafontaine
© 2017 Ryan A. St Laurent, Daniel Herbin, Carlos G. C. Mielke.
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, Herbin D, Mielke CGC (2017) Revision of the genus Reinmara Schaus, 1928 (Lepidoptera, Mimallonoidea, Mimallonidae) with the descriptions of four new species from South America. ZooKeys 677: 97-129. https://doi.org/10.3897/zookeys.677.12435
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Abstract
The mimallonid genus Reinmara Schaus, 1928 is revised. The three previously described species, R. enthona (Schaus, 1905), R. minasa Schaus, 1928, and R. wolfei Herbin & C. Mielke, 2014 are redescribed and the females of each are described and figured for the first time. Additionally, we describe four new species, two Andean: R. andensis sp. n. and R. occidentalis sp. n., and two Brazilian: R. atlantica sp. n. and R. ignea sp. n.. The new species R. ignea and R. atlantica are likely of conservation concern due to their rarity in collections and their apparent endemism to an endangered biome, the Brazilian Atlantic Forest.
Keywords
Bolivia, Brazil, Ecuador, Peru, Taxonomy
Introduction
The type species of Reinmara Schaus, 1928, R. enthona (Schaus, 1905), was originally described in Cicinnus Blanchard, 1852. Cicinnus was, and to some extent, still is a sort of catchall category subsuming many taxa of uncertain phylogenetic position. Later,
Since
Methods
Dissections were performed as in
Specimens from the following collections were examined:
CDH Coll. Daniel Herbin, Garidech, France
CEIOC Entomological Collection of the Oswaldo Cruz Institute, Rio de Janeiro, Rio de Janeiro, Brazil
CGCM Coll. Carlos G. C. Mielke, Curitiba, Paraná, Brazil
CPC Coll. Philippe Collet, Caen, France
ISEZ The Institute of Systematics and Evolution of Animals of the Polish Academy of Sciences, Kraków, Poland
MGCL McGuire Center for Lepidoptera & Biodiversity, Gainesville, Florida, USA
MWM Museum Witt, Munich, Germany
VOB Becker Collection, Camacã, Bahia, Brazil
Figures were manipulated with Adobe Photoshop CS4 (
We used DNA barcoding to help distinguish the similar species Reinmara enthona from R. andensis sp. n. Our barcoding protocol used a standardized short sequence of DNA as a species-level character (
The symbol ‡ is used in the text to represent unavailable names in the text (
Results and discussion
Reinmara
Type species
Cicinnus enthona Schaus, 1905;
Diagnosis
Reinmara can be recognized by the usual contrast between medial and submarginal areas due to diffuse, lighter coloration medially, this coloration, combined with the straight forewing postmedial line, notched tornus, and elongated, slightly falcate forewings (males) distinguish this genus from most other Mimallonidae. The morphologically most similar genus, Trogoptera Herrich-Schäffer, [1856], has more rectangular forewings and often displays more earthen tones (except in T. semililacea (Dognin, 1916) which is similar in color to some Reinmara but can be recognized by extremely long saccular extensions in the male genitalia). Genitalia of Trogoptera are very similar to those of Reinmara, but the fused gnathos is mesally extended by a singular structure, and is not distally separated as is the same structure in Reinmara.
Description
Male.Head: Pale beige to brown, eyes very large, occupying more than two-thirds area of head; antenna pale brown, tan, bipectinate to tip with distal fifth of pectinations much shorter; labial palpus reduced, not extending beyond frons, three segmented, second segment roughly half length of first, third segment reduced, barely visible; vestigial proboscis present. Thorax: Coloration usually as for head but with additional, often pink, shading. Legs: Coloration as for thorax, vestiture thick, long; tibial spurs narrow, very sharp, basal half covered in scales. Forewing dorsum: Forewing length: 12.0–23.5 mm, wingspan: 33–43 mm. Triangular, outer margin concave to varying degrees mesally; tornus usually strongly notched, apex may appear somewhat falcate in species with prominently concave outer margin. Ground color various shades of brown, sparsely scattered with dark brown, tiny petiolate scales usually present. Ante- and medial areas nearly always with pale pink or almost silvery scales throughout, submarginal area generally appearing darker than medial area. Presence of antemedial line variable, dark postmedial line preapical, well defined. Discal mark always present as pale splotch, with darker central region faint or very prominent. Fringe coloration variable. Forewing ventrum: Similar to dorsum but appearing browner overall due to absence of well-defined ante- and medial pink shading, though some pink shading may be present, especially submarginally. Antemedial line absent, postmedial line reduced to traces in all but R. ignea sp. n., discal mark more prominent, darker than on forewing dorsum. Hindwing dorsum: Shape more rounded, outer margin convex except for when notch present on anterior margin, patterning as for forewing dorsum, but antemedial line absent, discal mark and postmedial line usually weakly defined. Hindwing ventrum: Following same pattern as forewing ventrum. Frenulum a single bristle. Venation: Typical of Mimallonidae, but Rs3 + Rs4 quite long stalked. Abdomen: Coloration usually as for thorax, but browner, with coppery luster in fresh specimens, fading to pale beige in older material. Vestiture thick, long, distal tip of abdomen with elongated, dark-brown tipped scales. Genitalia: Vinculum ovoid, circumscribing a complex diaphragm with four setae-filled sacks, from a ventral perspective: upper two sacs much smaller and outwardly everted with long outwardly extended setae, lower two sacks larger (bottom right sack the largest of the four), lower sacks not outwardly everted, setae of lower sacks extended outward from within sacks. Uncus simple, broad, truncated to varying degrees distally, appearing beak-like laterally. Gnathos robust, proximally rectangular or rounded, with broad, dual mesal extensions that are fused together near base but bifurcate as fingerlike tips distally. Valves broad, short, rounded apically, sacculus accentuated as slight fold with both blunt and sharp projections near distal most portion of fold, length of sharp projection usually asymmetrical when comparing sacculus of both valves. Juxta partially fused to ventrum of phallus, basally juxta as widened lip where affixed to vinculum. Base of relatively small phallus narrower than distal portion, distal half of phallus variable in shape. Vesica very small, globular, with singular, long narrower extension. Female.Head: As for male but slightly broader; antenna dentate with very small pectinations along entire length of flagellum, except in R. ignea sp. n. where antenna more similar to that of male, but with smaller pectinations. Thorax: As for male. Legs: As for male. Forewing dorsum: Forewing length: 12–23 mm, wingspan: 27–43 mm. As for male but much broader, margin convex except for just below apex; tornus strongly notched. Coloration and patterning usually as for male, but see R. ignea sp. n. Forewing ventrum: Similar to dorsum but appearing browner overall due to absence of well-defined ante- and medial pink shading. Antemedial line absent, postmedial line usually reduced to traces, discal mark more prominent, darker than on forewing dorsum. Hindwing dorsum: Similar to forewing dorsum, but notch present on anterior margin, patterning as for forewing dorsum, but antemedial line absent, discal mark and postmedial line usually weakly defined. Hindwing ventrum: Following same pattern as forewing ventrum. Frenulum as multiple bristles. Abdomen: Similar to that of males but more robust overall. Genitalia: Stout, usually robust; tergite VIII forms smooth, posteriorly directed tongue-like extension, VIII heavily sclerotized laterally forming curving plate, which extends outward encircling papillae anales. Apophyses anteriores roughly half-length or equal to that of apophyses posteriores. Lamella ante- and postvaginalis converge as a wide, bowl-like structure. Ductus bursae short, narrow. Corpus bursae small in comparison to robust, heavily sclerotized remainder of genitalia, either bag-like or elongated. Papillae anales broad, rounded, covered in long, fine setae.
Remarks
The genus Reinmara is broadly distributed in South America. Prior to this study very little was known about the genus and females were unknown.
Unlike most genera of Mimallonidae studied by us in recent years, Reinmara have very homogenous male genitalia with only minor differences between externally distinct species (for example R. enthona and R. minasa), so we relied heavily on external characters, considering habitat specialization and endemism to specific habitats/biomes as seen in other mimallonid genera, as well as in one case COI barcoding, to differentiate species. We also recognize the close similarity in wing shape and male genitalia morphology between Reinmara and Trogoptera, but maintain them as separate, valid genera pending ongoing phylogenetic work.
Key to species of Reinmara
| 1 | Size in both sexes moderate (forewing length: >16 mm), forewing antemedial line very faint, if present at all; ventrally, postmedial line on all wings weakly defined, outwardly curved, usually interrupted by wing veins; forewing apex not falcate or if so, weak, blunt. Male genitalia: phallus cylindrical, weakly curved (for example Figs |
2 |
| – | Size in both sexes relatively small (12 mm [♂], 12–16 mm [♀]), forewing antemedial line present, not faint; ventrally postmedial line on all wings essentially as well defined and following the same pattern as on dorsum; forewing apex sharply acute, falcate. Male genitalia: phallus thin, strongly curved, and hook-like in shape (Fig. |
R. ignea sp. n. |
| 2 | Male: Forewing postmedial line not outwardly lined with black suffusion from tornus to apex. Female: Forewing postmedial line not inwardly lined with light pink-gray suffusion, medial area pink suffused | 3 |
| – | Male: Forewing postmedial line outwardly lined with black suffusion from tornus to apex. Female: Forewing postmedial line inwardly lined with light pink-gray suffusion, medial area largely displaying the light brown ground color, not suffused with pink | R. minasa |
| 3 | Forewing with deep notch at tornus, eastern slopes of the Andes mountains | 4 |
| – | Forewing smooth at tornus, notch absent, west of Andes | R. occidentalis sp. n. |
| 4 | Phallus mostly cylindrical in shape, not distinctly broadened distally, found in the Amazon rainforest, moderate elevations of the Andes mountains, or from the Brazilian Atlantic Forest | 5 |
| – | Phallus distinctly broadened distally, endemic to the Cerrado of Brazil and adjacent regions of Bolivia | R. wolfei |
| 5 | Forewing postmedial line notched toward costa at intersection with Rs4; forewing narrowed apically, slightly falcate, distributed in the Amazon rainforest and Brazilian Atlantic Forest | 6 |
| – | Forewing postmedial line not notched toward costa at intersection with Rs4; forewing not noticeably narrowed apically, though if somewhat truncated, submarginal area still broader than any other Reinmara species; endemic to eastern slopes of Andes mountains | R. andensis sp. n. |
| 6 | Setae-filled diaphragmal sacks of male genitalia well developed, extending into body cavity well beyond vincular ring. Pinkish gray suffusion generally broadly distributed in medial area of forewing. Broadly distributed in the Amazon rainforest | R. enthona |
| – | Diaphragmal sacks of male genitalia half the size of those in R. enthona, sacks hardly extending into body beyond vinculum. Gray suffusion of medial area restricted to apical confluence of postmedial line with costa. Endemic to Brazilian Atlantic Forest, so far known only from Espírito Santo | R. atlantica sp. n. |
Reinmara enthona
Cicinnus enthona Schaus, 1905: 325–326
Reinmara
enthona
;
Reinmara
enthona
;
Reinmara
enthona
;
Reinmara
enthona
;
Type material
Holotype, ♂. FRENCH GUIANA: St. Jean, Maroni, F. Guiana/ [Holo]Type, No. 8888, U.S.N.M./
Additional material examined
(67 ♂, 3 ♀ total) SURINAME: 2 ♂, Moengo, Boven Cottica River: 26.V.1927, Cornell Univ. Lot 760, Sub 79, 80, St Laurent diss.: 10-25-15:2 (
Diagnosis
Reinmara enthona, the type species of the genus Reinmara, is recognizable by the extensive suffusion of pinkish gray in the medial area. It is very similar to the following two species, but of the three species, R. enthona has the most extensive rosy medial suffusion, and a narrow submarginal area with quite falcate forewings (like R. atlantica sp. n., but unlike R. andensis sp. n.). The genitalia are intermediate in size between those of R. andensis and R. atlantica. The large diaphragmal sacks of R. enthona are similar to, but still smaller than those of R. andensis, whereas the same sacks of R. atlantica are about 50% smaller.
Description
Male.Head: As for genus, but light brown in color. Thorax: Coloration as for head. Legs: Coloration as for thorax, vestiture thick, long. Forewing dorsum: Forewing length: 16–22 mm, avg.: 19 mm, wingspan: 36–43 mm, n=16. Triangular, outer margin concave below apex; tornus notched, apex usually somewhat falcate. Ground color light brown to rich chocolate brown, very sparsely scattered with tiny, dark-brown, petiolate scales. Ante- and medial areas lighter brown than darker brown submarginal area, though in some specimens medial area may be very dark brown with less suffusion of grayish pink, lighter pinkish-gray scales present throughout medial area, including near costa on outer edge of postmedial line. Antemedial line almost nonexistent. Discal spot dark ovoid mark, surrounded by pale-gray scales, darker central area variable in expanse. Fringe coloration lighter brown than wing margin. Forewing ventrum: Similar to dorsum but more homogenously brown overall with very obvious black splotch at costa where postmedial line meets it, covering of dark petiolate scales may be much more extensive than on dorsum. Antemedial line absent, postmedial line reduced to traces. Hindwing dorsum: Notch on anterior margin weak, patterning as for forewing dorsum, but antemedial line absent, discal mark and postmedial line weakly defined. Hindwing ventrum: Following same pattern as forewing ventrum but traces of postmedial line outwardly bent mesally. Abdomen: Coloration as for thorax. Genitalia: (Fig.
Distribution
(Fig.
Remarks
Considering the expansive distribution of R. enthona, this name potentially includes several cryptic species. This section of the genus Reinmara warrants future investigation, especially on the lower and moderate elevations of the eastern Andes Mountains. We call attention to specimens from moderate elevations in Peru (MWM) and those from about 1400 m in Ecuador (MGCL) which could be R. enthona, R. andensis sp. n., or additional taxa. See remarks of R. andensis sp. n. for further discussion on this matter.
Reinmara atlantica sp. n.
Type material
Holotype, ♂. BRAZIL: Espírito Santo: BRASIL: ES, Linhares. 40 m, 05–09.iv.1992, V.O.Becker Col/ Col. BECKER 82019/
Paratypes
(3 ♂ total) BRAZIL: Espírito Santo: 2 ♂, same data and Becker number as the holotype,
Diagnosis
Reinmara atlantica is very similar to R. enthona but is darker brown, usually slightly smaller, and has narrower forewings. Also, the light gray medial suffusions are mostly restricted to area along the postmedial line, especially near the costa, and are not present throughout the medial region as in R. enthona. The postmedial line is slightly angled toward the costa at Rs4 in R. atlantica, not interrupted there in R. enthona. The genitalia can be recognized by the narrower valves and smaller gnathos extensions relative to the whole of the genitalia. Perhaps the most reliable character differentiating these two species is the reduced size of all four diaphragmal sacs, especially noticeable in the lower right sac which is very reduced in comparison to that of R. enthona, and hardly extends inward toward the body cavity, whereas this huge sac in R. enthona extends well into the body cavity past the vincular ring.
Description
Male.Head: As for genus, but light brown in color. Thorax: Coloration as for head. Legs: Coloration as for thorax, vestiture thick, long. Forewing dorsum: Forewing length: 19–20 mm, avg.: 19.7 mm, wingspan: 35–36 mm, n=3. Triangular, outer margin concave below apex; tornus notched, apex somewhat falcate. Ground color rich brown, very sparsely scattered with tiny, dark-brown, petiolate scales. Ante- and medial areas lighter brown than darker brown submarginal area, lighter gray scales present near costa on both sides of postmedial line, but more expansive on inner side with narrow strip of suffusion scales along postmedial line, fading before anterior wing margin, small patch of light-gray scales also present in antemedial area. Antemedial line almost nonexistent. Discal spot dark ovoid mark, surrounded by pale-gray scales. Fringe coloration lighter with nearly white trailing edge. Forewing ventrum: Similar to dorsum but more homogenously brown overall with very obvious black splotch at costa where postmedial line meets it. Antemedial line absent, postmedial reduced to traces. Hindwing dorsum: Notch on anterior margin weak, patterning as for forewing dorsum, but antemedial line absent, discal mark and postmedial line weakly defined. Hindwing ventrum: Following same pattern as forewing ventrum but traces of postmedial line outwardly bent mesally. Abdomen: Coloration as for thorax. Genitalia: (Fig.
Distribution
(Fig.
Etymology
This new species is named for the type locality, which is situated very near to the Atlantic coast of Brazil.
Remarks
Despite an abundance of Mimallonidae material from the Brazilian Atlantic Forest in collections visited during the course of this research (see list in Methods), the four specimens from Linhares were the only R. atlantica material located from this hyperdiverse biome. This species may be much more restricted within this biome than other species in the family that are also endemic to the Brazilian Atlantic Forest.
Reinmara andensis sp. n.
Type material
Holotype, ♂. BOLIVIA: BOLIVIE, N. Yungas, 1000–1800 m, Oct,nov,Dec,2008, Leg. local collector for R. Marx, Coll. D. Herbin/ genitalia prep. D. Herbin ref H. 1134/ HOLOTYPE male Reinmara andensis St Laurent, Herbin, & C. Mielke, 2017 [handwritten red label]/ (
Paratypes
(9 ♂ total) BOLIVIA: 1 ♂, same data as for holotype (CDH). La Paz: 1 ♂, Nor [North] Yungas, Road Caranavi-Coroico, ca. 100 km NE La Paz, ca. 16.2°S, 67.6°W, 1000–1800 m: V–VI.2009, R. Brechlin & F. Meister leg. (MWM). 1 ♂, Río Songo [recte Río Zongo], 750 m: ex-Coll. Fassl,
Specimens of uncertain identity hereby excluded from the type series
. ECUADOR: Napo: 1 ♂, 1 ♀, Wildsumaco Biol. Stat., E slope Andes Mtns, 0°40'17.2"S, 77°35'55.1"W, ~1400 m: 1–14.VIII.2016, Kawahara + Barber Labs et al. leg., DNA voucher numbers LEP-40632, 42829 (MGCL, molecular collection, barcoded). PERU: San Martín: 1 ♂, Mina de Sal, 1400 m: V.2007, Rainer Marx leg., Genitalia prep. No. 29.219 MWM (MWM). Huánuco: 1 ♂, Leoncio Prado, La Divisoria, 1600 m: 20.VI.1982, Charles F. Zeiger [leg.] (MGCL).
Diagnosis
Reinmara andensis is similar to R. enthona but larger, with broader wings and broader submarginal areas, which are more uniformly light brown. Medially the light gray scaling is reduced in comparison with R. enthona. The genitalia are very similar to those of R. enthona, but are overall somewhat larger, the gnathos extensions are shorter and phallus more tubular with a more protruding ventral distal lip in comparison with R. enthona. The lower right diaphragm sac is larger and more ovoid in shape in R. andensis, in R. enthona it is smaller and more spherical.
Description
Male.Head: As for genus, but light brown in color. Thorax: Coloration as for head. Legs: Coloration as for thorax, vestiture thick, long. Forewing dorsum: Forewing length: 18.5–20.0 mm, avg.: 19.2 mm, wingspan: 37–40 mm, n=5. Triangular, margin slightly concave below apex; tornus notched, apex hardly falcate. Ground color light orange-brown, very sparsely scattered with tiny, dark brown, petiolate scales. Ante- and medial areas appearing lighter brown than more uniformly orange-brown submarginal area due to suffusion of lighter gray scales medially, especially near costa and on inner side of postmedial line, in some specimens medial area may be very dark brown with less suffusion of grayish pink. Antemedial line almost nonexistent. Discal mark pale gray, ovoid, variously darkened at center. Fringe coloration lighter than wing margin with nearly white trailing edge. Forewing ventrum: Similar to dorsum but more homogenously brown overall due to reduction in paler gray shading. Antemedial line absent, postmedial line reduced to traces. Hindwing dorsum: Notch on anterior margin weak, patterning as for forewing dorsum, but antemedial line absent, discal mark and postmedial line weakly defined. Hindwing ventrum: Following same pattern as forewing ventrum but traces of postmedial line outwardly bent mesally. Abdomen: Coloration as for thorax. Genitalia: (Fig.
Distribution
(Fig.
Etymology
This new species is named for its Andean distribution.
Remarks
Additional material from MWM and MGCL from other localities in Peru besides those from the Puno region need verification due to the unreliability of the collector and/or unclear collecting data. We anticipate that this new species is more broadly distributed, but considering the close similarity to R. enthona and unavailability of recently collected Peruvian material, we restrict the type series of this species to include only those from northwestern Bolivia and adjacent southeast Peru. Although R. andensis is endemic to the eastern slopes of the Andes, it appears to be sympatric with R. enthona at the lower elevations in the inhabited range of R. andensis.
Due to the barcoding results (Fig.
In the
Reinmara occidentalis sp. n.
Psychocampa nocturna ‡ in Piñas 2007, fig. 215 ♂, nomen nudum
Type material
Holotype, ♂. ECUADOR: El Oro: ECUADOR, El Oro prov. 10km NW PIÑAS, 3°38'51"S, 79°45'52"W, 12.04.2012; H=750 m, leg. R. Brechlin & V. Sinyaev, Museum Witt/ Genitalpräparat Heterocera Nr. 29.218 Musuem WITT München/ HOLOTYPE male Reinmara occidentalis St Laurent, Herbin, & C. Mielke, 2017 [handwritten red label]/ (MWM). Type locality: Ecuador, El Oro, 10 km NW of Piñas.
Paratype
ECUADOR: El Oro: 1 ♂, Road Piñas-Saracay, 3°39'52"S, 79°45'26"W, 800 m: 6.XII.2012, Sinyaev & Romanov, expedition Ron Brechlin leg., genitalia prep. 30.813 (MWM).
Diagnosis
Reinmara occidentalis is one of most obscurely colored species in the genus. This new species is recognizable by the lack of a well-defined notch on the forewing tornus, which is instead smooth, and by the dark brown submarginal coloration with an almost complete absence of gray/pink shading in the medial region. On the ventral surface of the wings, the postmedial line is more continuous and less intermittently notched than in R. enthona, R. atlantica, or R. andensis. The male genitalia are also unique in this species because the gnathos extensions are quite long and deeply divergent, and the phallus is somewhat twisted, noticeably bent, and broadened distally unlike any other in the genus. This species is so far the only Reinmara known from the western slopes of the Andes.
Description
Male.Head: As for genus, but dark brown in color. Thorax: Coloration as for head but slightly lighter brown. Legs: Coloration as for thorax, vestiture thick, long. Forewing dorsum: Forewing length: 22.5–23.5 mm, avg.: 23 mm, wingspan: 40–42 mm, n=2. Triangular, outer margin weakly concave below apex; tornus smooth, unnotched, apex somewhat falcate. Ground color brown, sparsely scattered with dark brown, tiny petiolate scales. Ante- and medial areas lighter brown than darker, chocolate brown submarginal area, lighter gray scales present near costa on both sides of postmedial line. Antemedial line light brown but darker than surrounding area, wavy. Discal mark ovoid, surrounded by pale gray scales. Fringe coloration lighter brown than submarginal area. Forewing ventrum: Similar to dorsum but more homogenously brown overall, pale gray shading more evident near apex and submarginally. Antemedial line absent, postmedial line as on dorsum but fainter. Hindwing dorsum: Anterior margin without notch, but edge flatter than mesal wing margin. Patterning as for forewing dorsum, but antemedial line absent, discal mark and postmedial line weakly defined. Hindwing ventrum: Following same pattern as forewing ventrum but postmedial line outwardly bent mesally. Abdomen: Coloration as for thorax. Genitalia: (Fig.
Distribution
(Fig.
Etymology
This new species is named for the western (occidentalis Latin) Andean distribution.
Remarks
We are only aware of two specimens of this new species. Although data is still lacking in regards to the extent of the distribution of R. occidentalis, the distribution as well as the external morphology of this species are quite distinct from all others in the genus.
A specimen that may represent this new species was figured (fig. 215) in the plates of Piñas (2007) with the unavailable name Psychocampa nocturna‡ Piñas assigned by the author. As per information available in Thöny and Piñas (2015,
Reinmara wolfei
Reinmara wolfei Herbin and Mielke, 2014: 144, figs ♂ 40, 41, 43
Type material
Holotype, ♂. BRAZIL: Maranhão: holotype, Reinmara wolfei HERBIN & MIELKE det./ Brésil, Maranhão, Feira Nova do Maranhão, Retiro, 480 m, 24/31-XII-2011, 07°00'31"S, 46°26'41"W, C. MIELKE leg./ DZ 15.713/ Genitalia prep. D. Herbin ref. H 953/ (
Additional material examined
(7 ♂, 4 ♀ total) BRAZIL: Maranhão: 1 ♂, Balsas, 8°38'S, 46°43'W, 525 m: Coll. EMBRAPA-CPAC No. 20907 (CPAC). Goiás: 1 ♂, 2 ♀, Leop. Bulhoes [Leopoldo de Bulhões]: XI.1935, III.1936, ex. coll. R. Spitz, H.R.P[earson] genitalia prep. 4184 [lost], NHMUK010354557, 010354558 (2 ♀,
Diagnosis
Reinmara wolfei is characterized by the small size, sandy, tan brown coloration, only very faint to absent paler shading medially, and a faint or absent discal mark on the hindwing ventrum. The phallus of R. wolfei is the shortest and broadest of the genus. The female genitalia are not overly distinct from those of R. enthona.
Description
Male.Head: As for genus, coloration brown, antenna coloration brown. Thorax: Coloration lighter brown than that of head. Legs: Coloration as for thorax. Forewing dorsum: Forewing length: 15–17 mm, avg.: 16.3 mm, wingspan: 30–36 mm, n=4. Triangular, outer margin concave, tornus weakly notched. Ground color sandy brown. Ante- and medial areas concolorous, submarginal area above tornus slightly darker brown than remainder of wing in fresh specimens, pale suffusion present on inner side of postmedial line near costa. Antemedial line faint brown, wavy, postmedial line slightly curved, usually thick, black. Discal mark weakly represented by pale splotch with darkened region centrally. Fringe coloration as for remainder of wing or slightly darker. Forewing ventrum: Similar to dorsum but pale suffusions most absent except near apex. Antemedial line absent, postmedial line reduced to wavy traces, discal mark more prominent, darker than on forewing dorsum. Hindwing dorsum: Notch on anterior margin weak, patterning as for forewing dorsum, but antemedial line absent, discal mark and postmedial line weakly defined. Hindwing ventrum: Following same pattern as forewing ventrum. Abdomen: Coloration as for thorax. Genitalia: (Fig.
Distribution
(Fig.
Remarks
We figure and describe the female of this species for the first time, as well as the first Bolivian record. Until now, this species was known only from the male holotype from Maranhão, Brazil. We note some minor external differences between the specimens from drier Cerrado and that of the wet Pantanal, such as the slightly smaller size and brighter coloration in the Pantanal specimen (Fig.
Reinmara minasa
Reinmara minasa Schaus, 1928: 655, fig. ♂ 88b
Reinmara
minasa
;
Reinmara
minasa
;
Reinmara minasa ; Herbin and Mielke, 2014
Type material
Holotype, ♂. BRAZIL: Minas Gerais: Passa Quatro, Sul de Minas [SE of Minas Gerais], S.O. Brasilien, Jos. Zikán [leg.]/ [Holo]Typus/ No. [illegible] 6, 19-I-22/ Reinmara minasa Schaus type/ (
Additional specimens examined
(39 ♂, 2 ♀ total) BRAZIL: Espírito Santo: 1 ♂, No additional data, St Laurent diss.: 5-15-16:1 (
Diagnosis
This unique species of Reinmara can be recognized by the black suffusion along the entire length of the forewing postmedial line in males, which reaches the apex, darkening it. In both males and females there is a well-defined, narrow, pale pink suffusion along the postmedial line (outside of the black suffusion of the males, which is absent in females), leaving the remainder of the medial area mostly clear of pale pink suffusions. The male genitalia is recognizable by the uniformly narrow phallus with a usually distinctly backward splayed distal ventral tip, the uncus is quite broad. Among the species for which the female is known, R. minasa female genitalia is characterized by the largest dorsal projection of the tergite VIII as well as by the robustness of the lateral plates below the papillae anales.
Description
Male.Head: As for genus, coloration light brown. Thorax: Coloration as for head but with pale pink scales present on prothoracic collar and base of wings. Legs: Coloration as for thorax, but with additional, dark petiolate scales sparsely scattered amongst vestiture, tarsus yellower. Forewing dorsum: Forewing length: 16.5–21.0 mm, avg.: 18.1 mm, wingspan: 33.0–42.5 mm, n=9. Acutely triangular, narrow, outer margin concave; tornus deeply notched nearly until postmedial line, apex somewhat falcate. Ground color brown, very sparsely scattered with dark brown, tiny petiolate scales. Antemedial area with pale pink hue, medial area displaying narrow strip of ground color between pink hue of antemedial area and inner pink suffusion of postmedial line, submarginal area darker brown than medial area with pale gray lunule-like marking on margin and strong, black suffusion on outer edge of postmedial line, black suffusion becoming widest and more diffuse near tornus, extending along entire postmedial line to apex. Antemedial line hardly distinguishable but present as outwardly bent brown wave, postmedial line nearly straight. Discal mark variable from pale pink splotch with little to no black scales in center to almost entirely covered by black scales. Fringe coloration nearly white with darker scales at wing vein intersections. Forewing ventrum: As for genus but pale pink scales along postmedial line broadly scattered, postmedial line as on dorsum straight, but only fainter, black suffusion replaces lunule-like submarginal shape of dorsum. Hindwing dorsum: Notch on anterior margin weak, patterning as for forewing dorsum, but antemedial line absent, discal mark nearly always absent, pale suffusion submarginally similar to forewing lunule-like area. Hindwing ventrum: Following same pattern as forewing ventrum but postmedial line wavier, discal mark present, pale pink suffusion widely expanded throughout medial and submarginal areas. Abdomen: As for genus. Genitalia: (Fig.
Distribution
(Fig.
Remarks
Until now, very little has been reported on this species. We figure and describe the female of R. minasa for the first time.
Reinmara ignea sp. n.
Type material
Holotype, ♀. BRAZIL: Santa Catarina: BRAZIL – SC, São Bento do Sul, Rio Natal, 550 m., (no date). I. Rank leg./ 20.982 Col. C. Mielke [dissection number equivalent]/ HOLOTYPE female Reinmara ignea St Laurent, Herbin, C. Mielke, 2017 [handwritten red label]/ (
Paratypes
(1 ♂, 1 ♀ total) BRAZIL: Santa Catarina: 1 ♂, São Bento do Sul, Rio Vermelho, 968 m: 26.II.1973, A. & J. Razowski leg., St Laurent diss.: 5-6-16:1 (ISEZ). Rio de Janeiro: 1 ♀, Nova Friburgo, 1100 m: 21.I.1998, V.O. Becker leg., ex. Coll. Becker 112810, St Laurent diss.: 2-29-16:1 (
Diagnosis
This unique species cannot be confused with any other Mimallonidae. Reinmara ignea is the smallest species of Reinmara, bearing little outward resemblance to others of the genus. The tiny size, sharply acute and falcate forewings, thick postmedial and antemedial lines, narrow and curving phallus, are just the most immediately recognizable characters enabling the identification of this new species. We also note that this is the only species of Reinmara for which the female has bipectinate antennae like the male (albeit smaller overall), not dentate as in other female Reinmara.
Description
Male.Head: As for genus but coloration pale beige, antenna coloration pale brown due to scaling, but much darker brown beneath scales, vestigial proboscis not visible. Thorax: Coloration as for head. Legs: Coloration as for thorax, vestiture homogenously colored. Forewing dorsum: Forewing length: 12 mm, wingspan: 24 mm, n=1. Triangular, outer margin concave; tornus weakly notched, apex falcate. Ground color light orange-brown, speckling of tiny petiolate scales. Ante- and medial areas concolorous, darker brown than submarginal area, submarginal area much lighter orange-brown, appearing nearly yellow, faint pale lunule-like marking along margin below apex. Antemedial line defined, dark brown, slightly outwardly bowed, postmedial line also dark brown, slightly wider than antemedial line, barely curved. Discal mark as pale splotch, with obscured, darker central region. Fringe not well preserved. Forewing ventrum: Compared to forewing dorsum, more subdued tan brown, homogenous across all areas of wing, antemedial line absent, postmedial line as for dorsum, petiolate scaling heavier, especially antemedially, discal mark dark brown streak. Hindwing dorsum: Shape more rounded than forewing, outer margin convex except straight anterior margin, patterning as for forewing dorsum but both ante- and medial areas lighter, more similar to submarginal area in coloration, antemedial line absent, postmedial line as for forewing dorsum, well defined, discal mark present but weakly as pale streak. Hindwing ventrum: Following same pattern as forewing ventrum. Abdomen: As for genus. Genitalia: (Fig.
Distribution
(Fig.
Etymology
This new species is named for its fiery (ignea Latin) coloration, reminiscent of burning embers.
Remarks
Until the first author dissected the single male of this new species, proper generic placement was not clear to us, and we had originally considered R. ignea as belonging to an undescribed genus. Despite the outward uniqueness of both sexes, the genitalia of both sexes display characters fundamental to the diagnosis of the genus Reinmara. In the male, the structure and shape of the valves, the broad, mesally fused but distally separated gnathos, and balloon-like setae-filled sacs extending inward into the body cavity from the diaphragm are all typical of Reinmara, the gnathos character precluding R. ignea from placement in the related Trogoptera. Female genitalia are similar to those of other species of Reinmara, but the tergite VIII extension is particularly weakly sclerotized and thin (though present). We also note that this is the only species in the genus for which the female antennae are similar (bipectinate) to those of the male, just smaller, as in most mimallonid genera, not dentate as in the females of R. enthona, R. wolfei, and R. minasa.
We note minor difference in maculation of the two female specimens of R. ignea (compare Figs
This species and R. atlantica may very well be of conservation concern due to the present state of fragmentation of the biome to which they are endemic (
Two additional female specimens were located in the collection of Ivo Rank, collector of the holotype, but they are not included in the type series.
Reinmara enthona adults, a dorsal b ventral. 1 Holotype ♂, French Guiana, St. Jean du Maroni (
Reinmara adults, a dorsal b ventral. 8 R. atlantica Holotype ♂, Brazil, Espírito Santo, Linhares, 40 m (
Reinmara wolfei adults, a dorsal b ventral. 13 Holotype ♂, Brazil, Maranhão, Feira Nova do Maranhão, 480 m [image originally published by Antenor, reused with permission] (
Reinmara adults, a dorsal b ventral. 17 R. minasa Holotype ♂, Brazil, Minas Gerais, Passa Quatro (
Reinmara male genitalia, a ventral b lateral c phallus lateral. 25 R. andensis Holotype, Bolivia, N. Yungas, 1000–1800 m, D. Herbin genitalia prep. H. 1134 (
Reinmara male genitalia, a ventral b lateral c phallus lateral. 28 R. wolfei, Brazil, Distrito Federal, Planaltina, 1000 m, C. Mielke genitalia prep. 6.812 (CPAC) 29 R. minasa, Brazil, Espírito Santo, St Laurent diss.: 5-15-16:1 (
Reinmara female genitalia, a ventral b lateral c dorsal. 31 R. enthona, French Guiana, Kaw Rd., PK 37.5 + 2, 200 m, D. Herbin genitalia prep. H. 1103 (CDH) 32 R. wolfei, Brazil, Distrito Federal, Planaltina, 1000 m, C. Mielke genitalia prep. 6.879 (CPAC) 33 R. minasa, Brazil, São Paulo, Santo Antônio do Pinhal, Eugênio Lefèvre, 1200 m, C. Mielke genitalia prep. 28.071 (
Reinmara ignea female genitalia, a ventral b lateral c dorsal. 34 Holotype ♀, Brazil, São Bento do Sul, Rio Natal, 550 m, C. Mielke genitalia prep. 20.982 (
Acknowledgements
The following individuals and their respective institutions were vital to this research in enabling us to examine and study Reinmara material: Courtney Richenbacher and David Grimaldi (
References
- Adobe (2008) Photoshop CS4: version 11.0.
- Barcode of Life [or BOLD or BOLDSYSTEMS] (2013) Advancing species identification and discovery by providing an integrated environment for the assembly and application of DNA barcodes. Barcode of life data systems. http://www.barcodinglife.org or www. boldsystems.org [2/6/2017]
- Becker VO (1996) Mimallonidae. In: Atlas of Neotropical Lepidoptera, Checklist. Part 4B. Drepanoidea, Bombycoidea, Sphingoidea. Association for Tropical Lepidoptera & Scientific Publishers, Gainesville, Florida, 17–19.
- Fletcher DS, Nye IWB (1982) Bombycoidea, Castnioidea, Cossoidea, Mimallonoidea, Sesioidea, Sphingoidea, Zygaenoidea. In: Nye IWB (Ed.) The generic names of moths of the world. Trustees of the BMNH, London, 192 pp.
- Gaede M (1931) Pars 50: Mimallonidae. In: Strand E (Ed.) , Lepidopterorum Catalogus.W. Junk, Berlin, 2–19.
- Hebert PDN, Cywinska A, Ball SL, deWaard JR (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society B: Biological Sciences, 270, 313–321. https://doi.org/10.1098/rspb.2002.2218
- Herbin D, Mielke C (2014) Preliminary list of Mimallonidae from Feira Nova do Maranhão, Maranhão, northern Brazil with description of some new species. Antenor 1(2): 130–152.
- Kimura M (1980) A simple method for estimating evolutionary rate of base substitutions through comparative studies of nucleotide sequences. Journal of molecular Evolution, 16: 111–120. https://doi.org/10.1007/BF01731581
- Kristensen NP (2003) 4. Skeleton and muscles: adults. In: Kristensen NP (Ed.) Band 4: Arthropoda, 2 Hälfte: Insecta, Lepidoptera, Moths and Butterflies, Teilband/Part 36, Vol 2: Morphology, Physiology, and Development.Walter de Gruyter, Berlin, Boston, 39–131. https://doi.org/10.1515/9783110893724.39
- Lafontaine JD (2004) The moths of North America. Noctuoidea: Noctuidae (part). Noctuinae (part Agrotini). Fasc. 27.1. In: Dominick R, Ferguson D, Franclemont J, Hodges R, Munroe E (Eds) Wedge Entomological Research Foundation, Washington, DC, 385 pp.
- Piñas Rubio F (2007) Mariposas del Ecuador, Vol. 19. Compania Jesus, Quito, Ecuador.
- Ribeiro MC, Metzger JP, Martensen AC, Ponzoni FJ, Hirota MM (2009) The Brazilian Atlantic Forest: How much is left, and how is the remaining forest distributed? Implications for conservation. Biological Conservation 142: 1141–1153. https://doi.org/10.1016/j.biocon.2009.02.021
- Saitou N, Nei M (1987) The neighbor-joining method: A new method for reconstructing phylogenetic trees. Molecular Biology and Evolution 4: 406–425.
- Schaus W (1905) Descriptions of new South American moths. Proceedings of the United States National Museum 29: 179–345. https://doi.org/10.5479/si.00963801.1420.179
- Schaus W (1928) Familie Mimallonidae. In: Seitz A (Ed.) Die Gross Schmetterlinge der Erde.6. Die amerikanischen Spinner und Schwärmer. A. Kernen, Stuttgart, 635–672.
- Stoeckle MY, Hebert PDN (2008) Barcode of Life. Scientific American, 299: 82–88.
- Sukumaran J, Knowles LL (2017) Multispecies coalescent delimits structure, not species. Proceedings of the National Academy of Sciences 114(7): 1607–1612. https://doi.org/10.1073/pnas.1607921114
- Tamura K, Stecher G, Peterson D, Filipski A, Kumar S (2013) MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution 30(12): 2725–2729. https://doi.org/10.1093/molbev/mst197
- Thöny H, Piñas F (2015) Remarks, Corrections, Comments and Additions to Vol. 23 and 25 of the “Mariposas del Ecuador” on the Genus Gonodonta Hübner, 1818. Lepidoptera Research 25(1): 15–21.
- Thöny H, Piñas Rubio F (2017) Bemerkungen, Korrekturen, Kommentare und Nachträge zu „Mariposas del Ecuador“ von Piñas Rubio, 2005, Vol. 22 & 25, Noctuidae, Plusiinae 3. Beitrag zur Heterocera – Fauna von Ecuador (Lepidoptera, Noctuoidae, Noctuidae, Plusiinae). Entomofauna, Zeitschrift Für Entomologie 38(11): 213–232.