Roelofa hegewischi ( Druce, 1887 )

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 : 529-535

Roelofa hegewischi ( Druce, 1887)

( Figs 42 , 44–46, 50–52 , 53–57 )

Perophora hegewischi Druce, 1887 , Tab. 24, fig. 3

Roelofa hegewishi ; Schaus 1928: 640, fig. 87c, misspelling

Roelofa hegewischi ; Gaede 1931

Roelofa hegewischi ; Becker 1996

Roelofa hegewischi ; St Laurent and Kawahara 2019

Roelofa hegewischi ; St Laurent et al. 2020

Type material: HOLOYPE ♀. MÉXICO: Hegewischi Druce / Typus/ México. Hegewisch/ Coll. Staudinger./ Coll. Staudinger R. 916./ Type. Sp. figured./ Origin./ Perophora hegewischi type Druce/ ( MNHU, examined) . No paratypes.

Additional material examined: (95 ♂, 23 ♀ total) México: Estado de México: 1 ♂, 1 ♀, Malinalco: VI.1959, T. Escalante leg., UF FLMNH MGCL 1032506, 1032533 (MGCL). 3 ♂, Zacualpan: no date (1 ♂), 1918 (1 ♂), V.1922 (1 ♂), Joicey Coll. Brit. Mus. 1925-157, NHMUK010890574–010890576 (NHMUK). 3 ♂, Zacualpan: “II 14” [II.1914?]; “14”; VII.1913, 172 (MNHU). Chiapas: 1 ♂, San Cristobal de las Casas: 3.V.1973, H.L. King leg., UF FLMNH MGCL 1032472 (MGCL). 1 ♂, Hwy 195, Jitotol–Rayon, Cabañas Siempre Verde env., 17°08’34.67’’N, 92°53’01.14’’W, 1695 m: 16.V.2015, S. Naumann & B. Wenczel leg., St Laurent dissection: 8-17-17:3 (MGCL). 1 ♂, 11 km N of Ososingo, 1400 m: 18.IX.1993, B. Kelly leg. (CRAS). 1 ♂, Santa Rosa Comitan: VIII.1948, T. Escalante leg., UF FLMNH MGCL 1032657 (MGCL). 1 ♂, Hwy 190, 5 km E Rizo del Oro, 16°28’04.91’’N, 94°01’44.44’’W, 820 m: 14.V.2015, S. Naumann & B. Wenczel leg., St Laurent dissection and barcode: 8-17-17:4 (MGCL). 1 ♂, 28 mi. west Cintalapa: 9.IV.1962, F.D. Parker, L.A. Stange leg. (BME). Oaxaca: 1 ♂, 27 km 9° NE Huatulco, near Finca Monte-Carlo, 15°59.6’N, 96°06.3’W, 890 m: 26–31.VIII.2011, V. Siniaev & O. Romanov leg., expedition Dr. R. Brechlin (MWM). Querétaro: 6 ♀, Jalpan [de Serra]: 21.VIII.1988 (2 ♀), 1 ♀ with St Lau- rent dissection: 3-5-20:1, 2.IX.1991 (2 ♀), 4.IX.1991 (2 ♀), J. Adams leg. (CRAS). San Luis Potosí: 1 ♂, Ciudad del Maiz à El Naranjo, Piste de Maguey del Oriente km 1.1, N22°29.268’, W99°25.070’, 2829 ft: 7.VIII.2003, J. Haxaire leg. (CDH). 1 ♂, Ciudad del Maiz à El Naranjo, km 6.8, 22°26.660’N, 99°34.064’W, 1330 m: 27.VII.2000, J. Haxaire & O. Paquit leg. (CDH). Veracruz: 1 ♂, Misantla: VI.18, coll. Joh. Laue (ZSM). 2 ♂, 2 ♀, Jalapa [ recte Xalapa?]: 19.IV.13, 3.V.13 (2 ♂, ZSM); Collection of John T. Mason, Donated 1918 (1 ♀, DMNS); Collec- tion Wm. Schaus, USNM-Mimal: 1288 (1 ♀, USNM). 5 ♂, Huatuxco [ recte Huatusco], Rothschild Bequest BM 1939-1, NHMUK010890577–010890580, NHMUK010890583, genitalia prep. NHMUK010402322 (for specimen NHMUK010890583) (NHMUK). 1♀, No additional data, Rothschild Bequest BM 1939-1, NHMUK010890584, genitalia prep. NHMUK010402327 (NHMUK). No state data: 1 ♂, Collection BrklynMus, USNM-Mimal: 1290 (USNM). BELIZE: Cayo: 4 ♂, Mtn. Pine Ridge, 1000’ Falls: 23.V.1990 (1 ♂) 28.VI.1990 (2 ♂), 29.VI.1990 (1 ♂), Linwood C. Dow leg., UF FLMNH MGCL 1032505, 1032596, 1032639, MGCL Acc. #2015-16 L. Dow (MGCL). GUATEMALA: Alta Verapaz: 3 ♂, 1 ♀, San Cristobal [Verapaz] 4000 ft: V.1916, J.D. Norton leg., Ex. Coll. HJ Elwes 1920, Joicey Coll. Brit. Mus. 1925-157, HJ Turner BM 1949-586, Ex. AE Gibbs Coll H. Reynolds 1921-132, NHMUK010890569–010890572 (NHMUK). 2 ♀, San Cristobal [Verapaz?]: 1917, Rothschild Bequest BM 1939-1, NHMUK010890567, 010890568 (NHMUK). Baja Verapaz: 4 ♂, Quetzal Res., “Los Ranchitos,” 1680–1750 m: 10–15.VI.2007, J.B. Heppner leg., UF FLMNH MGCL 1032568 (MGCL). 1 ♂, Quetzal Res., “Los Ranchitos,” 1680 m: 27–30.VI.2012, J.B. Heppner leg., UF FLMNH MGCL 1032642 (MGCL). 1 ♂, Quetzal Res., “Los Ranchitos,” 1680 m: 3–6.VI.2017, J.B. Heppner and E. Fuller leg. (MGCL). 3 ♂, Biotopo del Quetzal, 15°12.952’N, 90°13.175’W, 1720 m: 23.V.2007, BC-Her 2757, 2761, D. Herbin & M. L. Montagnani leg. (CDH). El Progreso: 2 ♂, Cerro Pi[ñ]alon, Bosque Pino [pine forest], 15.07298°, -89.94833°, 2219 m: 16–18.V.2010, J. Monzón, B. Sutton, G. Steck, P. Skelley, St Laurent dissection: 8-17-17:9 (CJM). 2 ♂, Cerro Piñalon, Estación de campo Hector, Canteno, N15°05.037’, W89°56.563’, 2555 m: 27–28.IV.2017, J. Monzón, A. Mendez, & S. Naumann leg. (MGCL). Guatemala: 1 ♂, 1 ♀, Guatemala City: Rodriguez leg., Rothschild Bequest BM 1939-1, God- man-Salvin Coll. 98.40., NHMUK010890566, 010890573 (NHMUK). San Marcos: 2 ♂, San Miguel Ixtahuacán, Mina Marlin , 15.225717°, -91.698268°, 2015 m: 2.V.2011, Camposeco & Monzón leg., St Laurent dissection: 8-17-17:10 (CJM). Zacapa: 5 ♂, San Lorenzo, El Naranjo, 15.07329°, -89.68482°, 1616 m: 30.V.2009, J. Monzón & B. Sutton (1 ♂, CJM); 22–24.V.2010, J. Monzón, B. Sutton, G. Steck, P. Skelley leg., St Laurent dissection: 8- 17-17:5 (2 ♂, CJM); 30.V.2009, BC-Her 4297, J. Monzón leg. (2 ♂, CDH). 2 ♂, 1 ♀, 5 km SE La Union, Finca los Chorros, 14.942529°, -89.275854°, 1474 m: 12–15.V.2009, BC-Her 4184, Monzón and Camposeco leg. (CDH). 1 ♂, 3 km SE La Union, Finca los Chorros, N14°56.560’, W89°16.554’, 1443 m: 29–30.IV.2017, J. Monzón & S. Naumann leg. (MGCL). 2 ♂, SE La Union, 14°57.156’N, 89°16.689’W, 1421 m: 15.V.2007, BC-Her 2759, D. Herbin & M. L. Montagnani leg. (CDH). 3 ♂, N of San Lorenzo, 15.1023°, -89.6720°, 1902 m: 1.VI.2016, Peter Landolt leg. (CPL). 3 ♂, Sierra de las Minas, 12 km N of San Lorenzo, 3433 ft: 4–6.V.2006 (CPL). 13 ♂, 3 ♀, Sierra de las Minas, Cerro Monos, 15.11593°, -89.68541°, 2250 m: V.2008 (BC-Her 3681, 3683 [genitalia prep. D. Herbin ref H. 815], 3684) (6 ♂, 1 ♀), 1–3.VI.2009 [BC-Her 4104, 4105] (2 ♂, 1 ♀), 20.V.2010 (3 ♂), J. Monzón leg. (11 ♂, 2 ♀ total in CDH); 1–3.VI.2009, Monzón & B. Sutton leg., St Laurent dissections: 8-17-17:7 (1 ♂, 1 ♀); 19–21.V.2010, J. Monzón, B. Sutton, G. Steck, P. Skelley [leg.], St Laurent dissection: 8-17-17:6 (1 ♂) (2 ♂, 1 ♀ total in CJM). 3 ♂, Sierra de las Minas, N of Rio Hondo, E of San Lorenzo, Cerro Monos, N15°06.971’, W89°40.674’, 2243 m: 1–2.V.2017, J. Monzón & S. Naumann leg. (MGCL). 1 ♀, Sierra de las Minas, “Santa Cruz”, “Marble Quarry Rd”, NE of Teculutan, N15°03.224’, W89°40.737’, 955 m: 21.V.2006, MV/bl, P.J. Landolt leg. (CDH). 1 ♂, Sierra de las Minas, 15°04’33’’N, 89°41’33’’W, 1640 m: 13.X.2014, Barbut et al. leg. (CDH). No department data: 2 ♂, 3 ♀, G. Brückner S. G. (MNHU). 1 ♂, Georg Brückner (ZSM). 1 ♂ (RBINS). COSTA RICA: Cartago: 2 ♂, Orosi, Vulkan Jrazu [Irazú Volcano], 1200 m: Coll. Fassl. Teplitz, Joicey Coll. Brit. Mus. 1925-157, NHMUK010890581, 010890582, genitalia prep. NHMUK010402324 (NHMUK). PANAMA: Chiriqui: 1 ♂, Alto Quiel, 1650 m: IV.2010 (CDH). 1 ♂, Mt. Totumas Lodge, nr. Volcan, 1900 m: 24–30.V.2014, J.B. Heppner leg., UF FLMNH MGCL 1032617 (MGCL). 7 ♂, near Volcan town, 8.885329°, -082.683527°, 1900–2050 m: V–VI.2018, A. Kozlov & Yu. Kovaleva leg. (MGCL).

Additional records from the literature/ online resources: México: Chiapas: 1 ♂, La Trinitaria, 16.117605°, -91.674685°, 3.V.2016 (iNaturalist, observed by aleturkmen). HONDURAS: Francisco Morazán: Sex unknown, Uyuca hill, Uyuca Biological Reserve (RBU) ( Usedo 2016). 2 ♂, Centro Zamorano, Reserva Biológica Monte Uyuca: 20.X.2019, 3.VI.2020 (iNaturalist, observed by Eric van den Berghe).

Diagnosis. Compared to all previous species, R. hegewischi can be recognized by the nearly uniform pale, salmon pink ground color, sharply falcate forewings, and double postmedial lines. Genitalia of R. hegewischi and R. monzoni are distinguished from those of other Roelofa by the combination of narrow gnathos arms which terminate in spatulate tips, usually with a distinct ventral projection; simple, yet broad basal-valvae upward turned lobes, and particularly elongated, well developed juxtal projections that are usually strongly attached to the dorsum of the phallus and which are longer than the phallus itself. Roelofa hegewischi can be distinguished from R. monzoni by the slightly different shade of pink coloration between both species (see Figs 44–49 ) and the complete to nearly complete absence of a secondary postmedial line in R. monzoni .

Description. Male. Head: Coloration salmon pink, structure as for genus; antenna coloration as for genus, antennal structure as in R. olivia and R. maricia , though smaller overall, basally bipectinate distally finely serrate, appearing filiform. Thorax: Coloration deep salmon pink. Legs: Vestiture pale cream colored. Forewing dorsum: Forewing length: 15–23 mm, avg.: 19.2 mm, n = 41, wingspan: 29–46 mm. Triangular, outer margin smooth and concave below apex, becoming more convex mesally; apex falcate, usually sharply. Antemedial and medial ground color salmon pink, usually quite faded in preserved material, coloration may be deep pink in fresh specimens, overall lightly speckled with dark brown petiolate scales; postmedially pink coloration fades, becoming more khaki-brown. Costal coloration only slightly darker than remainder of wing. Antemedial line essentially absent, dual postmedial line well-defined, inner line browner in color, outer line less complete, darker, blackish brown, inner line meets apical streak after perpendicular angle toward costa at Rs 3. Discal mark a pale pink band along outer margin of discal cell, usually very faint. Forewing ventrum: Coloration and patterning similar to forewing dorsum, but antemedial line, outer postmedial line, and discal spot absent; medially coloration may be gray. Hindwing dorsum: Following similar patterning to forewing dorsum, antemedial line and discal spot absent. Hindwing ventrum: Following same pattern as forewing ventrum though markings may be almost nonexistent. Abdomen: Extending beyond anal hindwing margin, coloration as for thorax. Vestiture thick, compact, distal tip of abdomen with pair of elongated, dark-brown scale tufts that heavily contrast against remainder of abdomen and may contribute to about one quarter overall length of abdomen in living and well-preserved specimens. Genitalia: ( Fig. 53–57 ) n = 9. Vinculum ovoid, ventrally projected as small spine, diaphragm with dense region of elongated, but partially deciduous setae. Uncus simple, triangular, apically narrowed. Gnathos robust, proximally ovoid, with broad, dual mesal arms that are fused together basally, mesal arms narrow and fingerlike along their length but always distally flattened, usually with a slight downward projection at terminus, length of distal arms longer than that of proximal portion of gnathos. Valvae narrow, rounded apically, mostly simple except for inner mesal base, which is extended and modified as upturned fingerlike protrusion on each valva, which meet above phallus, protrusions not more heavily sclerotized than valvae; base of valvae notched on inner margin. Juxta partially fused to ventrum of phallus, and dorsally attached to diaphragm with pair of membranous processes that are longer than distal portion of phallus, these processes either remain attached to diaphragm or weakly attached to phallus upon removal of phallus. Phallus simple, pistol-shaped with curving coecum curved downward below, basal quarter of phallus smooth, distally lateral region of phallus covered by patch of small, semideciduous spines. Vesica elongate, bag-like. Female. Head: As for male but antenna finely serrate, appearing almost entirely filiform. Thorax: As for male. Legs: As for male. Forewing dorsum: Forewing length: 20–27 mm, avg.: 24 mm, n = 11, wingspan: 42–45 mm. As for male, but broader overall, apex less falcate. Forewing ventrum: Similar to forewing dorsum, but coloration paler overall. Hindwing dorsum: Following similar patterning to forewing dorsum, but discal spot absent. Hindwing ventrum: Following same pattern as forewing ventrum. Abdomen: As for male but more robust overall, distal tip of abdomen lacking paired elongated scale tufts, but distal tip of abdomen with singular darker scaled tuft. Genitalia: ( Fig. 42 ) n = 2. VIII a thickly sclerotized ring, continuous around circumference of segment, lamella antevaginalis a simple sclerotized band, lamella postvaginalis not sclerotized, dorsally VIII posteriorly smoothly curved along posterior margin with medial crease mesally. Apophyses anteriores vestigial, almost nonexistent, apophyses posteriores thickly sclerotized, somewhat irregular in width along length. Ductus bursae thick, tubular, corpus bursae bag-like, ovoid; together not much longer than remainder of genitalia. Papillae anales with setae shorter than overall length of papillae anales basally and much longer apically. Both examined dissections contained setae and potentially juxtal projections in ductus bursae, which are assumed to be derived from male.

Distribution. ( Fig. 59 ) Moderate to high elevations (862 up to 2555 m) in México and Central America, see remarks regarding localities lower than 800 m. Most records are from mountainous regions of southern México and Guatemala, though records exist from Belize, Honduras, Costa Rica, and Panama. We expect this species in western Nicaragua as well, but there has been less sampling in this country historically (pers. obs.).

Remarks. Roelofa hegewischi was described from a single female specimen, which can be inferred from Druce (1887) who said “[it is] the only one I have seen of this species.” The specific type locality of this species in México is uncertain, as originally stated by Druce “it is without any exact Mexican locality, but most likely came from the southern part of that country.” Based on the records available to us, we agree that the type likely came from southern México.

Roelofa hegewischi has what appears to be a degree of elevational phenotypic plasticity since material from lower elevations such as those from Veracruz, Oaxaca, Chiapas (e.g. 5 km E Rizo del Oro), San Luis Potosi, and Belize are phenotypically more similar to R. monzoni , mainly in the smaller size, narrower wing shape, more curved postmedial line, and reduction (though it is still present) of the outer postmedial line. However, the coloration of all R. hegewischi populations seems mostly consistent, being of a different, darker shade of salmon pink compared to the lighter pink coloration of R. monzoni . Dense genetic sampling of various populations of R. hegewischi and R. monzoni should be attempted in order to discern whether several cryptic taxa exist. Though our COI barcoding of specimens throughout Guatemala suggests two clear lineages: R. hegewischi and R. monzoni described below. One lower elevation population that displays the slightly different phenotype from more typical R. hegewischi was also sequenced (RASBC817174 from Chiapas, México). This sample is sister to all R. hegewischi (with R. monzoni sister to this pair), suggesting that there may be additional cryptic taxa within our broader concept of R. hegewischi . However, due to limited genetic sampling and lack of consistent morphological characters, we identify all specimens as R. hegewischi , with only the unambiguously distinct (genetically and morphologically) R. monzoni from Finca Firmeza del Banco in Izabal, Guatemala considered diagnostically separable.

The apparently allopatric Costa Rican/Panamanian populations may be a reasonable candidate for further crypsis in the R. hegewischi / R. monzoni group. External appearance and genitalia (genitalia prep. NHMUK010402324, Fig. 55 ) do not suggest any obvious distinction of this population, though the genitalia are larger overall than those of most R. hegewischi . The gnathos arms of this particular specimen are more uniformly shaped at their termini than in other examined R. hegewischi and R. monzoni , though the gnathos is rather variable across various genitalia preparations of these taxa.

Figures

FIGURE 59. Map of collecting localities for Roelofa hegewischi and R. monzoni examined in this work.

FIGURES 40–42. Roelofa female genitalia, ventral view.40.R. olivia, Colombia, Cundinamarca, Guaxa, Bogotá, genitalia prep. NHMUK010402328 [originally published in St Laurent and Kawahara 2019, Zookeys, reused with permission] (NHMUK). 41. R. maricia, Brazil, Rio de Janeiro, Nova Friburgo, 600 m, St Laurent dissection: 3-5-20:2 [corpus bursae damaged and not figured] (USNM). 42. R. hegewischi, México, Querétaro, Jalpan de Serra, St Laurent dissection: 3-5-20:1 (CRAS). Scale bar = 1 mm.

FIGURES 44–52. Roelofa specimens, a = dorsal, b = ventral. 44. R. hegewischi, male, Guatemala, Zacapa, 3 km SE La Union, Finca los Chorros, 1443 m (MGCL). 45. R. hegewischi, male, Costa Rica, Cartago Orosi, Irazú Volcano, 1200 m, photo courtesy A. Giusti (NHMUK). 46. R. hegewischi male, México, Chiapas, Hwy 190, 5 km E. Rizo del Oro, 820 m [low elevation population] (MGCL). 47. R. monzoni holotype, male, Guatemala, Izabal, Morales, Finca Firmeza del Banco, Sierra de Caral, 600 m (UVGC). 48. R. monzoni paratype, male, Guatemala, Izabal, Firmeza, Rio Bobos, near Morales, 950 m (MGCL). 49. R. monzoni paratype, male, Guatemala, Izabal, Morales, Finca Firmeza del Banco, Sierra de Caral, 600 m (CDH). 50. R. hegewischi female, México, Querétaro, Jalpan de Serra (MGCL). 51. R. hegewischi, female, Guatemala, Zacapa, 5 km SE La Union, Finca los Chorros, 1474 m (CDH). 52. R. hegewischi holotype, female, México (MNHU). Scale bar = 1 cm.

FIGURES 53–58. Roelofa male genitalia. 53. R. hegewischi, México, Chiapas, Hwy 195, Jitotol – Rayon, Cabañas Siempre Verde, 1695 m, St Laurent dissection: 8-17-17:3 (MGCL). 54. R. hegewischi, Guatemala, Zacapa, Sierra de las Minas, San Lorenzo, Cerro Monos, 2250 m, St Laurent dissection: 8-17-17:6 (CJM). 55. R. hegewischi, Costa Rica, Cartago Orosi, Irazú Volcano, 1200 m, genitalia prep. NHMUK010402324 (NHMUK). 56. R. hegewischi, México, Veracruz, Huatusco, genitalia prep. NHMUK010402322 [low elevation population] (NHMUK). 57. R. hegewischi, México, Chiapas, Hwy 190, 5 km E. Rizo del Oro, 820 m, St Laurent dissection: 8-17-17 [low elevation population] (MGCL). 58. R. monzoni holotype, Guatemala, Izabal, Morales, Finca Firmeza del Banco, Sierra de Caral, 600 m, St Laurent dissection: 8-17-17:11 (UVGC). Scale bar = 1 mm.