Roelofa Schaus, 1928: 640

St Laurent, Ryan A., Herbin, Daniel & Kawahara, Akito Y., 2020, Revision of Roelofa Schaus, 1928 (Lepidoptera, Mimallonidae, Roelofinae) with a description of a new species, Zootaxa 4877 (3), pp. 505-538 : 507-509

Roelofa Schaus, 1928: 640

Type species. Perophora olivia Schaus, 1896 ; Schaus 1928: 640 by original designation.

Diagnosis. Among Mimallonidae , Roelofa does not closely resemble any other genus. The species belonging to Roelofa can be recognized by the straight or nearly straight postmedial line of the forewing which, upon reaching Rs 3, combines with a black streak reaching the wing apex. The genitalia are most similar to those of Menevia but display variously shaped lobes that extend outward from the mesal base of the valvae. The gnathos arms are generally species-specific in shape, but are always paired and elongated, nearly reaching the saccular edge of valve when in their natural position. The phallus is simple and is weakly connected to paired arms which extend dorsally over the phallus among a dense region of diaphragmal setae. These arms are not unlike the “juxtal processes” of St Laurent and Dombroskie (2016), but are not so strongly sclerotized nor affixed to the juxta. Spines on the phallus and certain diaphragmal setae in male genitalia are apparently deciduous due to them both being lost together in some dissections (and the ease at which they fall out while dissecting). Additionally, the vincular “tusks” of Menevia are absent in Roelofa . The female genitalia are robust structures, albeit rather nondescript, the VIII segment is heavily sclerotized and dorsally forms a posteriorly directed arch, the corpus bursae is thick and baglike, which together with the ductus bursae is not much longer than the remainder of the genitalia. The apophyses anteriores are absent or reduced to small lumps but the apophyses posteriores are present and usually very thick.

Description. Male. Head: Eyes very large, occupying more than two-thirds area of head; antenna pale brown, tan, basal half to two thirds bipectinate, distal remainder serrate or filiform; labial palpus highly reduced, hardly extending to edge of frons, apparently three segmented but individual segments extremely small, especially distalmost segment; vestigial proboscis present. Thorax: Coloration usually as for head, broad, clothed in thick vestiture. Legs: Coloration usually as for thorax, vestiture thick, long; tibial spurs small, often not visible due to them being hidden by longer tibial vestiture. Forewing dorsum: Forewing length: 10.5–26.0 mm, wingspan: 22.0–49.0 mm. Triangular, outer margin smooth or mesally convex; apex somewhat falcate, except in R. olivia which has blunter apices. Ground color various shades of brown or pink, usually sparsely scattered with dark brown, tiny petiolate scales. Ante- and medial areas concolorous, displaying ground color, submarginally lighter. Antemedial line usually absent, dark postmedial line singular or consisting of two parallel lines, line preapical, but connects to dark streak shading in apical region above Rs 3. Discal mark variable. Fringe coloration variable. Forewing ventrum: Essentially identical to forewing dorsum, only varying slightly in shading, with fainter maculation. Hindwing dorsum: Following similar patterning to forewing dorsum, but discal spot absent. Hindwing ventrum: Following same pattern as forewing ventrum, but postmedial line may be nearly absent. Frenulum apparently absent. Venation: Typical of Mimallonidae , but Rs 3 + Rs 4 quite long stalked with M1 arising as far as halfway along length of stalk of Rs 3 +Rs 4. Abdomen: Coloration as for thorax. Vestiture thick, long, distal tip of abdomen with elongated, dark-brown tipped scale tufts, tufts range in length from one quarter to one third length of abdomen. Genitalia: Vinculum ovoid, diaphragm with dense region of elongated setae. Uncus simple, distally blunt or narrowed. Gnathos robust, proximally rounded, with broad, dual mesal extensions that are fused together basally, mesal extensions variable in shape but always long, extending nearly to distal valvae margins. Valvae narrow, rounded apically, mostly simple except for inner mesal base, which is variously extended and modified, usually more heavily sclerotized than surrounding area. Juxta partially fused to ventrum of phallus. Phallus simple, pistol-shaped with curving coecum which may be angled perpendicularly below phallus, phallus may be distally spined. Vesica bag-like. Female. Head: As for male but antenna finely serrate, appearing almost entirely filiform except in R. elyanae where antennae as in male of that species. Thorax: As for male. Legs: As for male, though tibial spurs may be stouter. Forewing dorsum: Forewing length: 12.0–29.5 mm, wingspan: 22.5–54.0 mm. As for male, but broader overall with wider submarginal area. Forewing ventrum: Essentially identical to forewing dorsum, only varying slightly in shading, with fainter maculation. Hindwing dorsum: Following similar patterning to forewing dorsum, but discal spot absent. Hindwing ventrum: Following same pattern as forewing ventrum, but postmedial line may be nearly absent. Frenulum apparently absent. Venation: Similar to males, but M1 arises closer to discal cell, not far along stalk of Rs 3 +Rs 4. Abdomen: As for male but more robust overall, distal tip of abdomen lacking paired elongated scale tufts, but in well-preserved specimens singular, darkened tuft of scales may be present. Genitalia: Robust; VIII a thickly sclerotized ring, continuous around circumference of segment or with gap ventrally at juncture with ostium bursae and ductus bursae, lamella antevaginalis not defined, lamella postvaginalis, if present, a simple plate, dorsally VIII mesally protruded or notched, but always thick, arching posteriorly. Apophyses anteriores vestigial or absent, apophyses posteriores present, usually very thickly sclerotized, robust, irregular in width along length, thickest mesally, knob-like or pointed apically. Membranous region between VIII and papillae anales expansive, often covered in short, thick setae. Ductus bursae thick, tubular sometimes with proximal region much wider than distal region, corpus bursae bag-like, circular. Papillae anales large, well-sclerotized, with setae of variable length.

Remarks. Four species of Roelofa have been included in a recent phylogenomic study of Mimallonidae ( St Laurent et al. 2020) . According to the trees presented in St Laurent et al. (2020), R. elyanae is sister to the remain- der of the genus, with R. narga ( Schaus, 1905) a clade containing R. hegewischi ( Druce, 1887) and type species R. olivia . Roelofa maricia Schaus, 1928 , R. maera Schaus, 1913 , and R. monzoni were not included in that study, but they are morphologically very or relatively similar to the four species included in St Laurent et al. (2020). Therefore, we follow a phylogenetic arrangement in the treatment of Roelofa species below. Our COI tree ( Fig. 1 ) recovers the same relationships among the taxa that are shared with the phylogenomic study of St Laurent et al. (2020). Here we include R. monzoni , which is recovered sister to R. hegewischi . Although our COI tree lacks some species of Roelofa , this tree is used primarily to discern close relationships between sister species R. hegewischi and R. monzoni , as well as to document the relatively low genetic variability among widely separated R. olivia populations.

Roelofa is one of the most widely distributed mimallonid genera, but one which contains relatively few species. The genus is known from central México south through Central America into South America, occurring on both sides of the Andes, as well as throughout the Amazon, Cerrado (savannah), and Mata Atlântica (Atlantic Forest) biomes. Very little is known about the natural history of Roelofa , except for an anecdotal report by Dognin (1922) for R. olivia which purported colonial nest building and silken tunnels. Construction of silken tunnels is not known for any other Mimallonidae , and gregarious behavior is rare (but see Mesquita et al. 2010). Data pertaining to the life history of one other species, is also known, and is more typical of Mimallonidae larval natural history. See the remarks for these species below for additional information.

Considering the phylogenetic placement of Roelofa , sister to much of the diversity of Mimallonidae , including all genera except Zaphanta , Roelofa is certainly of interest from an evolutionary perspective. It will be important to investigate the natural history strategies of this genus, including host plants and larval behavior, in an effort to understand the broader evolutionary history of Mimallonidae .

Key to species of Roelofa

Due to the rather distinct differences in external morphology and patterning among the seven species of Roelofa , we offer a key to wing patterning as it is the simplest way to differentiate species in this genus.

1 Outwardly, postmedial line accompanied by prominent dots at intersections with wing veins.......................... 2

- Outwardly, postmedial line not accompanied by dots at intersections with wing veins, either with outer line paralleling postmedial line, or no external markings along postmedial line....................................................... 3

2 Pale yellow suffusion near the discal cell bleeds into the angle between M 3 and CuA 1, distribution in Brazilian Atlantic Forest only.................................................................................. R. maera stat. rev.

- Pale yellow suffusion near the discal cell does not bleed into angle between M 3 and CuA 1, present in the Amazon, foothills of Andes, northern Cerrado......................................................................... R. narga

3 Submarginal region golden yellow, contrasting against darker purple-brown medial and antemedial areas....... R. elyanae

- Coloration not as above................................................................................ 4

4 Hyaline discal spot present on forewing, distributed in southeastern Brazil................................ R. maricia

- Forewing lacks hyaline discal spot, distributed in Central America and the Andes Mountains......................... 5

5 Antemedial, medial, and submarginal areas mostly concolorous, not distinctly contrasting, Central American............ 6

- Lighter submarginal area starkly contrasting against dark brown medial and antemedial areas, Andean............ R. olivia

6 Forewing postmedial line doubled, with outer line paralleling darker, inner postmedial line................. R. hegewischi

- Forewing postmedial line singular (faint suffusion possible tornally, but not completely lining outer margin of postmedial line)................................................................................... R. monzoni sp. n.

Figures

FIGURE 1. Maximum likelihood (ML) tree of Roelofa inferred with IQ-TREE which was based on the mitochondrial COI “barcoding” gene, rooted to Zaphanta fraterna Schaus (Zaphantinae, sister lineage to all other Mimallonidae). Black circles indicate SH-aLRT ≥ 80 and UFBoot ≥ 95. The scale bar represents expected number of nucleotide substitutions per site. See Supplementary File 1 for all nodal support values.