Four new species of Symmerista Hübner, 1816 (Notodontidae, Nystaleinae) from Costa Rica

Abstract The genus Symmerista Hübner (Notodontidae, Nystaleinae) is reviewed for Costa Rica, based on 49 wild-caught specimens. Four species are newly described: Symmerista luisdiegogomezi Chacón, Symmerista inbioi Chacón, Symmerista minaei Chacón and Symmerista aura Chacón. All are from the cloud forests of the Talamanca moutain range, southern Costa Rica. Photographs of the adults, male and female genitalia, and barcodes are also provided. The species Symmerista tlotzin Schaus (1892) is removed from Symmerista and assigned to the genus Elymiotis Walker as a new combination.


Introduction
The family Notodontidae includes about 3500 described species worldwide (Weller 1992). Approximately 896 species in 127 genera have been recorded in Costa Rica (Chacón et al. 2013). The subfamily Nystaleinae contains approximately 300 species and is restricted almost entirely to the Neotropics (Miller 1991), with a few species extending as far north as Canada (Forbes 1948). In Costa Rica, 191 Nystaleinae species are known to occur.
Hübner (1821) established the genus Symmerista, which until now has contained 12 described species (Franclemont 1946;Thiaucourt 2007; Thiaucourt and Monzón rough, pubescent, ventral surface smooth with sparse pubescence; socii long, wide, pubescent at bases, narrow and flattened at apices, shape as in the figure; length of the phallus 3.1 mm, proximal part of phallus tube wide at the base, narrow in the middle, distal part of phallus tube robust, sclerotized, with a tubular lateral projection and rounded apex; proximal part of the vesica with a ventral scobinate patch, distal part of the vesica bulbous. Female genitalia: posterior apophyses longer and more slender than anterior apophyses; CB rounded and membranous, lacking a signum; DB wide and sclerotized; posterior margin of postvaginal plate sclerotized and emarginate. Description. Male (Figs 1,2,(5)(6)(7)(8). Head -Antenna bipectinate, dark brown, six terminal flagellomeres without rami, antennal shaft dark brown dorsally and beige ventrally; scape bearing a long tuft of beige scales; haustellum reduced to two small lobes; eye smooth, round, black; frons mostly dark brown and black with brownyellow scales; labial palpus porrect, dark brown ventrally, light brown dorsally; vertex brown yellow; patagium light brown. Thorax and abdomen -Tegula dark brown at base, a mix of cream and dark brown scales distally; mesoscutellum dark brown; thoracic pleuron from creamy white to dark brown; dorsal area of metathorax with black hair-like scales; legs mostly dark brown with cream-colored scales between segments; abdominal dorsum dirty beige, venter beige. Wings -Dorsal FW ground color dark brown, with reniform spot black; basal band light brown; M sinuous, light brown; postmedial band light brown; an irregular, thin cream-colored bar from reniform spot to apex; AD black; fringe dark brown with beige scales where veins touch termen;  . Genitalia ) -T8 wider than long, with two windows, anterior margin slightly concave, posterior margin sclerotized; St8 wide at base, narrowing posteriorly, posterior margin concave with tiny sclerotized teeth on edge, with a pair of short lateral projections, these sclerotized with blunt apices (Fig. 7); valva membranous, finely pubescent, costulae absent, sacculus smooth, its margin uniform, costa straight with a distal protuberance near apex; valva with two triangular spine-like processes, one at base, other at juxta near anellus; tegumen narrowed dorsally; uncus slightly concave, somewhat helmet shaped, dorsal surface rough, pubescent, ventral surface smooth with sparse pubescence; socii long, wide, pubescent at bases, narrow and flattened at apices, shape as in the figure; juxta ovoid; vinculum lightly sclerotized (Fig. 5); length of phallus 3.1 mm, proximal part of phallus tube wide at base, narrow in middle, distal part of phallus tube robust , sclerotized, with a tubular lateral projection and rounded apex; proximal part of vesica with a ventral scobinate patch, distal part of the vesica bulbous (Fig. 8).
Female (Figs 3,4,9) Similar to the male: Head -Antenna filiform, antennal shaft dark brown with cream scales, scape bearing a long tuft of cream scales; frons and vertex dark brown; haustellum vestigial; labial palpus dark brown. Thorax and abdomen -Mostly dark brown, tegula beige; dorsal area of metathorax with black hair-like scales; abdominal dorsum light brown, venter beige. Wings -Dorsal FW ground color dark brown; antemedial band beige, lined on both sides by sinuous dark brown lines; irregular white thin bar from apex to reniform spot; reniform spot black; postmedial band beige, subterminal line black; a light beige area between subterminal line and postmedial band of M 3 extending to tornus; fringe dark brown with beige scales located where veins touch termen. Dorsal HW dirty beige; postmedial line light brown; discal spot black; fringe dark brown with beige scales located where veins touch termen (Figs 3,4)  . Genitalia (Fig. 9) -Papillae anales slightly ovoid, setose; posterior apophyses longer and more slender than anterior apophyses; DB wide and sclerotized; CB rounded and membranous, lacking a signum; posterior margin of postvaginal plate sclerotized and emarginate.
Distribution and habitat. Symmerista luisdiegogomezi has been collected only between 2450 and 2600 m in the highland cloud forests dominated by Quercus trees in the foothills west of the Cordillera de Talamanca (Talamanca Mountain Range), southern Costa Rica (Fig. 49 Etymology. This species is dedicated to the Instituto Nacional de Biodiversidad (IN-Bio) in recognition of its 25 years of support for developing an understanding of the biodiversity of Costa Rica and exporting that understanding to the nation and the world.
Diagnosis. Symmerista inbioi differs from Symmerista luisdiegogomezi on: dorsal FW ground color light brown; from the reniform spot to the apex there is an uniform long, thin, white cream-colored band; fringe reddish brown; dorsal HW dirty beige. Male genitalia: St8 wide at base, anterior margin convex, posterior margin slightly irregular with a pair of very short projections, these sclerotized with blunt apices; valva membranous, saccular margin slightly jagged, costal margin smooth, with a distal protuberance near the apex; elongate finger-shaped process on the saccular margin at the internal base of the valve; uncus plate slightly concave, with papillae and setae on the dorsal and ventral edge; socii elongated, broad at base, narrow and flattened at the apex, with papillae and setae in the dorsal surface; length of the phallus 3.9 mm, proximal part of phallus curved at the base, slightly narrow in the middle, distal part of phallus tube sclerotized, narrow at the base, robust, irregular and wide at the end, with a tubular lateral projection with the distal nipple; proximal part of the vesica with a dorsal scobinate patch, distal part of the vesica bulbous. The genital armature of Symmerista inbioi is more robust and larger than Symmerista luisdiegogomezi.
Wings -Dorsal FW ground color light brown; with reniform spot black; from reniform spot to apex with a uniform, long, thin, creamy-white band; costal margin black until R 1 ; AD black; purple scales between terminal line and the AD; fringe reddish brown; dorsal HW dirty beige, fringe light brown (Figs 10, 11) . Male genitalia (Figs 12-16) -T8 rectangular, wider than long, posterior margin slightly sclerotized and serrated; St8 wide at base, anterior margin convex, posterior margin slightly irregular with a pair of very short projections, these sclerotized with blunt apices (Fig. 14); valva membranous, costulate absent, saccular margin slightly jagged, costal margin smooth, with a distal protuberance near apex; elongate finger-shaped process on saccular margin at internal base of valve; tegumen narrow at point of intersection of both arms, margins heavily sclerotized (Figs 12, 13); uncus plate concave, with papillae and setae on dorsal and ventral edge; socii elongated, broad at base, narrow and flattened at apex, with papillae and setae in dorsal surface (Figs 12,13,15); juxta heart shaped (Fig. 13); vinculum membranous, slightly sclerotized (Fig. 12); length of phallus 3.9 mm, proximal part of phallus curved at base, slightly narrow in middle, distal part of phallus tube sclerotized, narrow at base, robust, irregular and wide at end, with a tubular lateral projection with distal nipple; proximal part of vesica with a dorsal scobinate patch, distal part of vesica bulbous (Fig. 16).
Female: unknown Distribution and habitat. Symmerista inbioi has been collected only at elevations betwen 1250 and 2700 m in highland cloud forests of the Cordillera de Talamanca (Talamanca Mountain Range) (Fig. 49).
Remarks Etymology. This species is dedicated to the Ministerio del Ambiente y Energía (MI-NAE) of the government of Costa Rica in recognition of its 28 years of continuous and widespread support for the survival and conservation of the wild biodiversity of Costa Rica.
Diagnosis. Symmerista minaei differs from Symmerista luisdiegogomezi on: dorsal FW ground color beige and light brown, square mark creamy near the reniform spot; beige mark in the apex; fringe beige yellow; dorsal HW beige. Male genitalia: T8 anterior margin slightly concave, posterior margin finely serrated with a window in the center; St8 lateral margins wide at the base, narrow to posterior margin, anterior margin concave, slightly sclerotized with a short projection in the center, posterior margin with robust projections, highly sclerotized on each side, with blunt apices, a little dome in the middle of the posterior margin; length of the phallus 3.3 mm, proximal part of phallus tube wide at base, distal part of phallus tube robust, sclerotized, with a tubular lateral projection rounded apex with the distal nipple; proximal part of the vesica with a ventral scobinate patch, distal part of the vesica bulbous. Female genitalia: Anterior and posterior apophyses the same size, long an slender; DB sclerotized; CB rounded, membranous and pleated; posterior margin of postvaginal plate sclerotized, slightly irregular, inverted V-shape.
Thorax and abdomen -Tegula dark brown at base, a mix of dark brown and light brown scales distally; mesoscutellum beige and dark brown; thoracic pleuron from beige to dark brown; dorsal area of metathorax with black and dark brown hair-like scales; legs mostly dark brown with beige scales between segments; abdominal dorsum dirty beige, venter beige. Wings -dorsal FW ground color beige and light brown, with reniform spot black; basal band light brown; postmedial band light brown; a creamy-white square distal to reniform spot; AD black; fringe dark brown with beige scales where veins touch termen; beige mark in apex; dorsal HW beige; fringe beigeyellow (Figs 17, 18) (WL 17.20-19.21 mm). Male genitalia (Figs 21-24) -T8 wider than long, anterior margin slightly concave, posterior margin finely serrated with a window in center; St8 lateral margins wide at base, narrow to posterior margins, anterior margin convex, slightly sclerotized with a short projection in center, posterior margin concave with a pair of short projections, these sclerotized with blunt apices, a small setose dome in middle of posterior margin (Fig. 23); valva membranous, mildly pubescent, margin of sacculus slightly serrated, costa with straight margin with a distal protuberance close to apex; valva with a triangular spine-like process; tegumen narrowed dorsally; uncus plate slightly concave, somewhat helmet shaped, dorsal surface rough, pubescent, ventral surface smooth with sparse pubescence, socii long, wide, pubescent at bases, narrow and flattened at apex, form s-shaped; vinculum slightly sclerotized (Figs 21, 22); length of phallus 3.3 mm, proximal part of phallus tube wide at base, distal part of phallus tube robust , sclerotized, with a tubular lateral projection rounded apex with distal nipple; proximal part of vesica with a ventral scobinate patch, distal part of vesica bulbous (Fig. 24). Female (Figs 19, 20, 25) Head -Antenna simple, shaft dark brown with cream-colored scales, scape with a tuft of cream-colored scales; eyes naked; frons dark brown, vertex dark brown with groups of beige scales at base and between antennal bases; haustellum vestigial, labial palpus dark brown.
Thorax and abdomen -Generally dark brown, tegula beige, scales of thorax long and forked. Patagium and prothorax with beige and cream scales; mesothorax with scales cream, dark brown and beige; metathorax with a group of black hair-like scales along posterior margin; abdomen light brownish gray, abdominal apex with a thick group of beige scales.
Wings -dorsally FW ground color dark brown; antemedial beige band lined at both sides by sinuous dirty dark brown lines; reniform spot black; an irregular thin white to beige line extends from the apex to the reniform spot; postmedial line beige, lined on each side with dark brown; adterminal line black; light beige area between adterminal line and postmedial band from M3 to tornus; fringe dark brown with beige scales where veins touch termen; dorsal HW dirty beige, fringe beige (Figs 19, 20) (WL 19.63-21.18 mm). Female genitalia (Fig. 25) -Papillae anales mambranous with short, scattered setae with longer, inwardly-curved setae arising from base. Anterior and posterior apophyses of same size, long and slender; DB sclerotized; CB rounded, membranous and pleated; posterior margin of postvaginal plate sclerotized, slightly irregular, inverted V-shape. Distribution and habitat. Symmerista minaei has only been collected at elevations between 2400 and 2600 m in highland cloud forests of the Cordillera de Talamanca (Talamanca Mountain Range) (Fig. 49) Prov. Cartago, Turrialba, Tayutic, Moravia de Chirripo Shipiri, 9.837781-83.453639, 1000m, 10 May 1983.
Etymology. This species is dedicated to Isidro Chacón's daughter, Aura Chacón, for 25 years of understanding an absent father obsessed with his work.
Diagnosis. Dorsal FW ground color light gray; a white cream mark from the discal cell to the apex. Male genitalia: St8 anterior margin with a sclerotized triangular projection in the middle, posterior margin sclerotized, irregular, with a rectangular projection at the center bearing two highly sclerotic and sharply serrated structures; proximal part of phallus tube wide and short at the base, with a blade-like lateral projection, sharp at the distal apex; a small rounded projection and curve off the previous, distal part of phallus tube robust, sclerotized, with a small bulge on the ventral side; proximal part of the vesica with a ventral scobinate patch, distal part of the vesica bulbous. Female genitalia: papillae anales ovoid-triangular, setose; posterior apophyses longer and more slender than anterior apophyses; DB sclerotized; CB elongated, membranous and pleated, lacking a signum; margin distal of antevaginalis plate very sclerotized and lightly depressed at the center, concave to the sides, proximal margin convex.
The female of Symmerista aura differs from female of Symmerista meridionalis Thiaucourt, 2007 in the shape of the antivaginalis plate; the CB elongated, membranous and pleated, lacking a signum; margin distal of antevaginalis plate very sclerotized and lightly depressed at the center, concave to the sides, proximal margin convex. This is the description of the female genitalia of Symmerista meridionalis published by Thiaucourt in 2007 which mentiones the differences, "Female terminalia: distal edge of lamella postvaginalis slightly incurved; lamella antevaginalis rectangular, almost square; its distal margin strongly sclerotized; mouth of the ostium bursae oval, densely sclerotized under the margin; ductus bursae beyond the constriction under the ostium, forming a short funnel; bursa membranous inserted on the dorsal surface of the extremity of the duct; signum distinct." The male of Symmerista meridionalis is unknown.
Wings -Dorsal FW ground color light gray; mark white cream from the discal cell to apex; costal margin beige until R1; subterminal line black; fringe light gray; dorsal HW light gray, fringe cream (Figs 26, 27) (Figs 30-34) -T8 wider than long, posterior margin sclerotized and highly irregular; St8 anterior margin with a sclerotized triangular projection in middle, posterior margin sclerotized, irregular, with a rectangular projection at center bearing two highly sclerotic and sharply serrated structures (Fig. 34). Valvae membranous with saccular margin esclerotized, slightly irregular and setose; costal margin smooth, apex rounded, sclerotized and setose; tegumen with margins very sclerotized, narrowed where both arms intersect; uncus slightly concave plate, with papillae and setae in dorsal and ventral edge, with socii elongate, wide at base, narrowed and flattened at apex with papillae and setae in dorsal part (Figs 30,31,33); juxta heart shaped; vinculum membranous slightly sclerotized (Fig. 31); proximal part of phallus tube wide and short in base, with a blade-like lateral projection, sharp at distal apex and a small rounded curved projection, distal part of phallus tube robust, sclerotized, with a small bulge on ventral side; proximal part of vesica with a ventral scobinate patch, distal part of vesica bulbous (Fig. 32).

Male genitalia
Female (Figs 28,29,35) Similar to male: Head -Antenna filiform, antennal shaft light brown with cream scales, scape bearing a long tuft of cream scales; frons and vertex cream; haustellum vestigial; labial palpus mostly cream with black lateral scales. Thorax and abdomen -tegula light gray; mesoscutum dark brown anteriorly, light brown and cream posteriorly; mesoscutellum black and gray; thoracic pleuron beige; legs dark brown on outer surfaces, beige on inner ones; abdominal dorsum dark brown, venter beige. Wings -Dorsal FW ground color light gray; mark white cream from the discal cell to apex; costal margin beige until R1; subterminal line black; fringe light gray; dorsal HW light gray, fringe cream (Figs 28,29)  . Female genitalia (Fig. 35) -papillae anales ovoid triangular, setose; posterior apophyses longer and more slender than anterior apophyses; DB sclerotized; CB elongated, membranous and pleated, lacking signum; margin distal of antevaginalis plate very sclerotized and lightly depressed at center, concave to sides, proximal margin convex.
Distribution and habitat. In Costa Rica, Symmerista aura has been collected between 1000 to 1400 m on both slopes of the Cordillera de Talamanca (Talamanca Mountain Range) (Fig. 49) posed by three segments; haustellum is well developed, ocelli absent; eyes smooth, round. Thorax mostly gray, tegula gray, all scales of the thorax are long and forked; patagium and prothorax with beige and light brown scales. FW costa straight, outer margin almost straight; accessory cell present. Male genitalia -(Figs 41-43) valvae elongated, sclerotized and mildly setose; sacculus pleats highly developed; uncus thin and long, with apex acute and setose, each socius wide at the base with two thin projections, acute and setose; saccus acute (Figs 41); phallus robust, wide at the base with two single subterminal lateral bumps, vesica long but shorter than phallus, wide at the base, long and thin distally, caltrop cornuti (Fig. 42); T8 rectangular shorter than St8, lateral margins straight, anterior margin simple and membranous, posterior margin slightly concave; St8 wide at anterior margin, posterior margin with a single median depression and two sclerotized small projections (Fig. 43). Female genitalia (Figs 44, 45) -The papillae anales small, roughly ovoid with elongated dorsal setae; lateral procecess of postvaginal plate with apices acute and sclerotized; posterior apophyses short and thin; lamella postvaginalis rectangular, sclerotized; ostium sclerotized, wide, funnel shape; DB short; CB wide and long, ovoid with surface membranous slightly rough; signum brick shape with a rough surface. Thiaucourt (2007) "The species was described in the genus Edema, cited by DRU-CE (1898) in this genus, the species should be now in another genus." KIRBY (1892) lists Edema (Walker, 1855) as a junior synonym of Symmerista Hubner, [1821]. The adult lacks the apical white stripe, the male genitalia framework places the genus in the Nystaleini, but it differs from that of Symmerista (Plate II, Fig. 10). Male genitalia: uncus with acute and long apex; socci in brackets; valve costa without apical membranous area, pleats highly developed; exopenis with two single subterminal lateral bumps; beam cornuti extensions, obsolete; distal edge St 8 with a single median depression, two sclerotized thickenings near the distal edge of T8.
We propose that Symmerista tlotzin should be allocated to the genus Elymiotis Walker, 1857 for the following characteristics of the male genitalia: valvae elongated, sclerotized and setose; sacculus pleats highly developed; uncus thin and long, with apex acute and setose; socius wide at the base with two thin projections, acute and setose and caltrop cornuti.