Roelofa olivia ( Schaus, 1896 )

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 : 519-527

Roelofa olivia ( Schaus, 1896)

( Figs 23–27 , 31, 32, 34 , 36, 37 , 40 )

Perophora olivia Schaus, 1896: 52

Perophora olivia ; Dognin 1922

Roelofa olivia ; Schaus 1928, fig. 87c

Roelofa olivia ; Gaede 1931

Roelofa olivia ; Fletcher and Nye 1982

Roelofa olivia ; Becker 1996

Roelofa olivia ; St Laurent and Kawahara 2019

Roelofa olivia ; St Laurent et al. 2020

Type material: LECTOTYPE ♂. COLOMBIA: Colombia S America/ Collection Wm Schaus / Perophora olivia type Schaus/ USNM-Mimal: 1131/ Type No. 12560 U.S.N.M./ LECTOYPE Perophora olivia designated by St Laurent, Herbin and Kawahara/ ( USNM, examined) .

Additional material examined: (121 ♂, 13 ♀ total) COLOMBIA: Cundinamarca: 4 ♂, Chicaque Natu- ral Park, 4°36’21.15′ ′ N, 74°18’19.94′ ′ W, 2610 m: 7.V.2019, D. Herbin leg. (CDH). 2 ♂, 1 ♀, Parc Chicaque, 4°36’28.5′′ N 74°18’16.37′′ W, 2556 m: 2.V.2019, D. Herbin leg. BC-Her5247 (♂) (CDH). 2 ♂, 1 ♀, Bogotá, Pueblo Guasca (1 ♂, CUIC); collection Frank Johnson, USNM-Mimal: 1308, 1313 (1 ♂, 1 ♀, USNM). 3 ♂, Bogotá, Rothschild Bequest 1939-1, NHMUK010890533– 010890535 (NHMUK). 1 ♂, Bogotá: 4.VI.1918, Dognin Col- lection, USNM-Mimal: 2430 (USNM). 1 ♂, 5 ♀: Guaxa nr. Bogata [Bogotá]: V.1921, A Maria, Joicey Coll. Brit. Mus. 1925-157, NHMUK010606778, 010890536–010890541, genitalia prep. NHMUK010402328 [♀] (NHMUK). 1 ♂, 3 ♀, Paramos de Guasca, Bogotá: VII.1919, Dognin Collection, USNM-Mimal: 1306, 1307, 1309, 1310 (USNM). 1 ♂, Colombia, no other data (CUIC). 1 ♂, Colombia, other data illegible, Joicey Coll. Brit. Mus. 1925- 157, NHMUK010890537–010890538 (NHMUK). Boyaca: 2 ♂, Arcabuco, Vereda Rupavita, 5°44’28.73’’N, 73°24’01.73’’W, 2984 m: 3.V.2019, D. Herbin leg. (CDH). 3 ♂, Villa de Leiva [ recte Villa de Leyva], 17.VI.1996, 2500 m, R. Reitmaier leg. (MWM). 1 ♂, Santander, road Duitama-Charala, 5°58’13’’N, 73°10’07’’W, 2925 m: 15–17.III.2016, St Laurent dissection: 12-4-18:2 (MGCL). Valle del Cauca: 1 ♀, 30 km E from Buenaventura, 22–30.VI.1996, 200 m [elevation incorrect, locality questionable and not shown in map Fig. 43 ], R. Reitmaier leg. (MWM). Risaralda: 2 ♂, Termales de San Vicente, 04°51’18’’N, 75°31’46’’W, 2560 m: 18–21.I.2015, M. Márquez & J. Machado leg., St Laurent dissection: 8-17-17:1 [ex. A. Kozlov] (MGCL). No additional data: 1 ♂, Coll. Staudinger, 796 (MNHU). ECUADOR: Morona-Santiago: 1 ♂, Route Gualaceo-Mendez, km 54, 2300 m: 30.VII.1990, D. Herbin, J. Haxaire leg. (CDH). 1 ♂, Route Gualaceo-Mendez, km 58, 2300 m: 9.VIII.1988, D. Herbin, J. Haxaire leg. (CDH). 3 ♂, 9 km W Plan de Milagro to Gualaceo, 3°00’04’’S,78°30’49’’W, 2375 m: 15.II.2012 (2 ♂), 26.I.2012 (1 ♂), R. Brechlin & V. Siniaev leg. (MWM). 3 ♂, 30 km Road Plan de Milagro to Gualaceo, 3°00’21’’S, 78°38’28’’W, 2970 m: 1– 2.II.2012, R. Brechlin & V. Siniaev leg. (MWM). 4 ♂, 62 km road Rio Bamba-Macas, 2°12’40’’S, 78°23’51’’W, 2700 m: 27.III.2012, R. Brechlin & V. Siniaev leg. (MWM). 13 ♂, 34 km Road Plan de Milagro to Gualaceo, 3°00’13’’S, 78°38’46’’W, 3176 m: 19.XI.2011 (2 ♂), 30.I.2012 (10 ♂), 17.II.2012 (1 ♂), R. Brechlin & V. Siniaev leg. (MWM). 5 ♂, 1 ♀, 34 km Road Plan de Milagro to Gualaceo, 3°01’24’’S, 78°35’6’’W, 2157 m: 21.XI.2011 (5 ♂), 28.I.2012, genitalia prep. 35.538 (1 ♀), R. Brechlin & V. Sini- aev leg. (MWM). 1 ♂, Road Gualaceo-Plan de Milagro, 3°0’21’’S, 78°29’53’’ W, 2033 m: 22.XI.2011, V. Siniaev & O. Romanov leg. (MWM). 2 ♂, Road Gualaceo-Plan de Milagro, 3°0’42’’S, 78°36’19’’W, 2601 m: 17.XI.2011, 22.XI.2011, V. Siniaev & O. Romanov leg. (MWM). 1 ♂, Mendez, 2°44’37’’ S, 78°18’27’’ W, 480 m: 28.XI.2011, V. Siniaev leg., coll. Dr. Ronald Brechlin [this locality is highly questionable given its elevation, not shown in Fig. 43 ] (MWM). Loja: 2 ♂, Road Loja-Zamora, 3°59’19’’S, 79°08’39’’W, 2800 m: 27.II.2012, R. Brechlin & V. Sini- aev leg. (MWM). 3 ♂, Road Loja-Zamora, 3°58’45’’S, 79°08’28’’W, 2700 m: 22.II.2012, R. Brechlin & V. Siniaev leg. (MWM). 1 ♂, Road Loja-Zamora, 3°58’45’’S, 79°08’28’’W, 2714 m: 25.XI.2012, V. Siniaev & O. Romanov leg. (MWM). 2 ♂, 15 km E Loja to Zamora, 3°58’45’’S, 79°08’28’’W, 2700 m: 1.III.2011, 25.XI.2012, H. Kaech & R. Brechlin leg. (MWM). 1 ♂, ca. 10 km S Saraguro, Monte del Cerro Fierrurco Sector de Puzón, 3°41’32’’S, 79°17’42’’W, 3140 m: 28.II.2011, H. Kaech & R. Brechlin leg. (MWM). Napo/Cotopaxi: 2 ♂, Oriente, Salcedo/ Napo, Km 58, 3100 m: 16.IV.1991, BMNH(E)2008-107, NHMUK010588510, NHMUK010588526 (NHMUK). Napo: 1 ♂, Route Cosanga-Tena, km 7, 2350 m: 19–20.VII.1990, D.Herbin, J.Haxaire leg., genitalia prep. H1137 (CDH). 14 ♂, Papallacta, Rio San Pedro, 0°22’56’’S, 78°7’27’’W, 3010 m: 4.XI.2011 (6 ♂), 18.I.2012 (2 ♂), 22.III.2012 (6 ♂), R. Brechlin & V. Siniaev leg. (MWM). 12 ♂, Rio Papallacta, Cuyuja, 0°25’17’’S, 78°1’19’’W, 2525 m: 6.XI.2011, V. Siniaev & O. Romanov leg., genitalia prep. 35.545 (MWM). 1 ♂, Cordillera Guacamayos, 0°37’15’’S, 77°49’28’’W, 2181 m: 11.XI.2011, V. Siniaev & O. Romanov leg. (MWM). Pichincha: 1 ♂, Quito- Nanegalito, 37 km, 0°01’3’’N, 78°36’55’’W, 2094 m: V. Siniaev & O. Romanov leg. (MWM). 1 ♂, Guagua Pichin- cha, 0°06’20’’N, 78°34’19’’W, 3676 m: 21.X.2011, V. Siniaev & O. Romanov leg. (MWM). 3 ♂, Tandayapa, Bel- lavista Lodge, IX.2012, R. Beck leg., MWM genitalia prep. 35.546, St Laurent dissection: 8-17-17:2 (2 ♂ MWM, 1 ♂ MGCL). Carchi: 2 ♂, El Angel Ecol. Reserve, 0°46’14’’N, 78°03’27’’W, 2785 m: 9–11.XI.2012, Victor Sin- yaev leg., expedition Ron Brechlin (MWM). 5 ♂, El Angel Ecol. Reserve, road Tulcan-El Chical, 0°48’46’’N, 78°00’40’’W, 3300 m: 14.XI.2012, Victor Sinyaev leg., Expedition Ron Brechlin (MWM). 1 ♂, 70 km Road Tul- can-El Chical, 0°50’29’’N, 78°03’25’’W, 2440 m: 22.XI.2012, Sinyaev & Romanov leg., expedition Ron Brechlin (MWM). El Oro: 1 ♂, Road Piñas-Saracay, 3°38’57’’S, 79°45’17’’W, 850 m: 5.XII.2012, Sinyaev & Romanov leg., expedition Ron Brechlin, genitalia prep. 35.547 (MWM). Azuay: 1 ♂, 15 km E Gualaceo to Mendez, 2°56’9’’S, 78°42’45’’W, 2920 m: 27.II.2011, H. Kaech & R. Brechlin leg. (MWM). PERU: Amazonas: 2 ♂, Streck Bagua- Chica-Nazareth, 700–1100 m [elevation likely inaccurate]: X–XII.1998, R. Marx leg. (SMFL). Piura: 2 ♂: Cordil- lera de Guamani (W), Huancabamba prov., 30 km W Huancabamba, 5°20’S, 79°31’W, [WGS 84], 3250 m: I.2012, Arturo, Dr. Ronald Brechlin leg. (MWM). Junin: 1 ♂, 12 km Satipo, 11°16.1’S, 74°38.5’W, 660 m [the coordinates do not agree with the designation of 12 km from Satipo, and elevation is likely too low to be accurate for this species, this data point is not shown in Fig. 43 ]: 24.XII.2010, V. & S. Sinjaev(a) leg., coll. Dr. Ronald Brechlin, genitalia prep. 35.548 (MWM). Cusco: 1 ♂, 12 km SW Marcapata, 13°35.4’S, 70°57.9’W, 2850 m: 28–30.XI.2010, V. & S. Sinjaev(a) leg., coll. Dr. Ronald Brechlin, genitalia prep. 35.549 (MWM). No country data: 1 ♂, “ Perophora apicistriga ‡ Weym. (Weym.) Col. ”, “ olivia Schaus ”, 59 (MNHU).

Literature records: ECUADOR: Tungurahua: 1 ♀, Baños-Pelotero, 2800 m: 2.I.1998, G. Onore [leg.] (Pi- ñas 2004). Loja: 1 ♂, Sagraguro-Cerro Torre, 3500 m: 29.XII.1997, G. Onore [leg.] (Piñas 2004).

Diagnosis. This species distinguished by the large size, such that it is the largest in the genus, and is in fact one of the largest of the Mimallonidae considering the size of the female. The dark brown antemedial and medial areas contrast heavily against the pale pink-tan postmedial and submarginal areas and the yellowish brown discal spot. The male genitalia are very robust structures, with heavily sclerotized, rectangular gnathos arms and fingerlike inner mesal projections of the valvae. The phallus is devoid of spines, which are present in most other Roelofa . Roelofa olivia is an Andean species found at some of the highest elevations reported for Mimallonidae and can be confused with no other species where it is found. The most similar species, R. maricia , is found in southeastern Brazil, has narrower wings, acuter forewing apices, is overall lighter tan in color, and displays unique, circular hyaline discal spots. The male genitalia of R. maricia are less robust overall, with narrower gnathos arms and spinier lobe-like, rather than smooth and fingerlike, inner mesal valvae projections. Female genitalia are different as well, in R. olivia the posterior margin of the VIII tergite is not mesally projected, but rather flat and smooth. Ventrally the VIII segment is mostly membranous with only minor sclerotization on either side of the ostium bursae, this same region is well-sclerotized in R. maricia .

Description. Male. Head: Coloration rich dark brown, structure as for genus; antenna coloration as for genus, basal half of antenna bipectinate, pectinations becoming longer then dramatically shortening near halfway point along antenna, after which antenna finely serrate appearing nearly filiform. Thorax: Coloration rich deep chestnut brown to darker chocolate brown, appearing hoary due to presence of lighter brown or pinkish gray scales. Legs: Coloration as for thorax, vestiture thick, long. Forewing dorsum: Forewing length: 19.5–26.0 mm, avg.: 22.6 mm, n = 9, wingspan: 38.5–44.0 mm. Triangular, outer margin smooth and concave below apex; apex falcate. Antemedial and medial ground color a layering of light brown scales with dark brown scales speckling the wing surface, these areas overlaid with thin, hair-like light brown scales; postmedially very light cream darkening to pale khaki submarginally. Costa appearing lighter than remainder of wing due to pink scales. Antemedial line essentially absent, dark postmedial line consists of pair of dark brown preapical lines, faintly angled toward costa at Rs 3, postmedial line connects to elongate black streak of shading spanning from costa to Rs 3, reaching apex. Discal mark an ovoid yellow splotch which may be bisected by dark brown bar, actual width of splotch variable. Forewing ventrum: Essentially identical to forewing dorsum, discal spot less well-defined. Hindwing dorsum: Following similar patterning to forewing dorsum, discal spot absent. Hindwing ventrum: Following same pattern as forewing ventrum, but postmedial line may be faded and/or interrupted by veins. Abdomen: Robust, extending beyond anal hindwing margin, coloration as for thorax. Vestiture thick, long, distal tip of abdomen with pair of elongated, dark-brown tipped scale tufts, tufts may be as much as one third abdomen length. Genitalia: ( Figs 36, 37 ) n = 9. Vinculum ovoid, ventrally projected as small spine, diaphragm with dense region of elongated, partially deciduous setae. Uncus simple, triangular, apically narrowed. Gnathos robust, proximally ovoid, with broad, dual mesal extensions that are fused together basally, mesal extensions rectangular, may be somewhat splayed and flattened distally, length of distal arms equal to or 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 which meet above phallus, protrusions more heavily sclerotized than valvae and may be obfuscated by diaphragmal setae; base of valvae notched on inner margin. Juxta partially fused to ventrum of phallus, dorsally juxta with pair of very small (less than one eighth length of phallus) membranous processes attached to diaphragm. Phallus simple, pistol-shaped with curving coecum which may be angled perpendicularly below phallus or slightly backward from it, phallus distally smooth, without spines except for narrow strip which may be covered in fine, deciduous spines on both lateral sides (usually nearly entirely lost in most specimens). Vesica 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: 25.0–29.5 mm, avg.: 26.4 mm, n = 7, wingspan: 49–50 mm. As for male, but broader overall with wider submarginal area, only very weakly concave below apex. Forewing ventrum: Similar to forewing dorsum but maculation more diffuse, postmedial line particularly diffuse such that appearing as singular line band rather than parallel pair of lines. 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. Genitalia: ( Fig. 40 ) n = 2. Robust; VIII a thickly sclerotized ring, continuous around circumference of segment except for ventrally at juncture with ostium bursae and ductus bursae, no obvious lamella antevaginalis or lamella postvaginalis present due to membranous break in the sternal region, dorsally VIII mesally notched along anterior margin, posterior margin smooth. Apophyses anteriores vestigial, reduced to small knobs, apophyses posteriores very thickly sclerotized, robust, irregular in width along length, thickest mesally, knob-like distally. Membranous region between VIII and papillae anales covered in short, thick setae. Ductus bursae thick, tubular, corpus bursae bag-like, circular. Papillae anales large, well-sclerotized, with setae longer than overall length of papillae anales.

Distribution. ( Fig. 43 ) Roelofa olivia is an Andean species, encountered in Colombia, Ecuador, and Peru. Collecting bias in Ecuador results in most of the records being from this country. This species is likely more widely distributed in Colombia and Peru than our records indicate. Records exist from up to 3676 m in elevation, though most are from 2000–3000 m. Records from significantly lower elevations (200 m at Valle del Cauca, Colombia; 480 m at Mendez, Morona-Santiago, Ecuador; 660 m in Junín, Peru) are likely erroneous, which is particularly evident due to other conflicting information about some of these localities such as mismatch between given coordinates and locality designations (see material examined for specific notes about these records). The record from a moderate elevation of 850 m in El Oro, Ecuador appears to be reliable, however.

Biology. Dognin (1922) provided some anecdotal information pertaining to the life history of R. olivia . He reported that larvae of this species were found living in a colonial nest consisting of silken tunnels. Construction of silken tunnels has not been observed anywhere else in Mimallonidae . It is unclear whether later instar R. olivia larvae build individual cases as do all known Mimallonidae larvae (e.g. Stehr 1987, Lemaire and Minet 1998). Gregarious behavior is only otherwise known in Psychocampa Grote and Robinson, 1867 , from the common P. “ callipius ” (formally treated as belonging to Cicinnus Blanchard, 1852 , specific identity not yet confirmed) ( Mesquita et al. 2010).

Remarks. Prior to the present study, only the male genitalia of R. olivia had been figured ( St Laurent and Kawahara 2019, St Laurent et al. 2020). Genitalia are largely consistent across the wide distribution of R. olivia , although phallus width does seem to vary slightly. Dissections from eastern Andean localities in Ecuador revealed a slightly more broadened phallus than those from western Andean localities, and eastern Andean Peruvian localities displayed a more cylindrical phallus. Roelofa olivia can be encountered at very high elevations, well above 3000 m, with many records from central regions of the Andes mountain chain (that is, not only on eastern or western slopes) and so it is likely that gene flow across the eastern and western slopes of the Andes occurs at various points along the latitudinal gradient where this taxon occurs. Minor external variation can be seen within populations collected at single localities as per the several large series in the MWM, thus variation which is likewise slight between more widely separated populations, is not indicative of various cryptic species. Our barcoding results suggest some phylogenetic structure among R. olivia populations, however we were only able to sequence one western Andean specimen from Ecuador, a single specimen from the Cordillera Central in Colombia (Risaralda) and two from the Cordillera Oriental (Santander and Cundinamarca). The Ecuadorian specimen was found be sister to a clade containing the three Colombian specimens with strong support, but support was weak for the sister relationship between the Risaralda specimen and the two from Cundinamarca. Overall, genetic differentiation among R. olivia was lower than between sister species R. hegewischi and R. monzoni , for example. Therefore, we do not at this time recognize distinct species within R. olivia and suggest population-level phylogenetic analyses for future research in this group. Such analyses would be better suited for uncovering phylogenetic structure worthy of taxonomic recognition.

In the MNHU, there is a single specimen labeled “ Perophora apicistriga ‡ Weym. [er]” from Colombia. Schaus (1928) mentioned this specimen as well and could not find a reference for the name. We also could not locate any publication using this name, thus it is apparently unavailable, although the specimen in question is certainly R. olivia .

Roelofa olivia was described from an indeterminate number of specimens, but a single male labeled as the “type” in the USNM is the only known syntype. Therefore, we designate this specimen as the lectotype.

Figures

FIGURE 43. Map of collecting localities for Roelofa olivia and R. maricia examined in this work.

FIGURES 23–27. Roelofa olivia male specimens, a = dorsal, b = ventral. 23. Lectotype, Colombia (USNM). 24. Colombia, Cundinamarca, Chicaque Natural Park, 2610 m (CDH). 25. Ecuador, El Oro, road Piñas-Saracay, 850 m, photo courtesy N. Ignatev (MWM). 26. Peru, Cusco, 12 km SW Marcapata, 2850 m, photo courtesy N. Ignatev (MWM). 27. Ecuador, Napo, Salcedo/Napo, Km 58, 3100 m, photo courtesy A. Giusti (NHMUK). Scale bar = 1 cm.

FIGURES 31–35. Roelofa female specimens, a = dorsal, b = ventral. 31. R. olivia, Ecuador, Morona-Santiago, 34 km Road Plan de Milagro-Gualaceo, 2157 m, photo courtesy N. Ignatev (MWM). 32. R. olivia, Colombia, Guaxa, near Bogotá, photo courtesy A. Giusti [originally published in St Laurent and Kawahara 2019, Zookeys, reused with permission] (NHMUK). 33. R. maricia, Brazil, Minas Gerais, Poté, 500 m, photo courtesy N. Ignatev (MWM). 34. R. olivia, Colombia, Valle del Cauca, 30 km E from Buenaventura [elevation possibly incorrect] (MWM). 35. R. maricia, Brazil, Rio de Janeiro, Barreira, Teresópolis (DZUP). Scale bar = 1 cm.

FIGURES 36–39. Roelofa male genitalia, a = ventral, b = lateral, c = phallus. 36. R. olivia, Colombia, Santander, road Duitama- Charala, 2925 m, St Laurent dissection: 12-4-18:2 (MGCL). 37. R. olivia, Ecuador, Pichincha, Tandayapa, Bellavista Lodge, 2300 m, St Laurent dissection: 8-17-17:2 (MGCL). 38. R. maricia, Brazil, Rio de Janeiro, Barreira, Teresópolis, 350 m, genitalia prep. NHMUK010402325 [dorsal juxtal projections not attached to phallus] (NHMUK). 39. R. maricia, Brazil, Goiás, Leopoldo de Bulhões, genitalia prep. NHMUK010402326 (NHMUK). Scale bar = 1 mm.

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.