Cryptic species within cryptic moths: new species of Dunama Schaus (Notodontidae, Nystaleinae) in Costa Rica

Abstract Based on almost 1,700 recently reared and wild-collected specimens, the genus Dunama Schaus (Notodontidae, Nystaelinae) in Costa Rica is reviewed. Eight species are recorded of which seven are newly described: Dunama jessiehillae Chacón, Dunama jessiebarronae Chacón, Dunama janewaldronae Chacón, Dunama jessiebancroftae Chacón, Dunama janecoxae Chacón, Dunama biosise Chacón, Dunama indereci Chacón. Dunama angulinea Schaus is redescribed and associated with its correct genitalia. Dunama tuna (Schaus), previously listed as ocurring in Costa Rica, is restricted to Colombia. Most species are described through their distinctive CO1 barcodes, genitalia and life histories. Dunama adults and caterpillars, their foodplants, and their parasites in Area de Conservación Guanacaste (ACG) in northwestern Costa Rica are described where known. Many life history stages are illustrated.


Keywords
Dunama, Heliconia, Musa, Arecaceae, caterpillars, moths, inventory, DNA barcodes, tropical forest, Area de Conservacion Guanacaste, INBio introduction Schaus (1912) established the genus Dunama for a group of small, relatively drab, mottled and tree-bark patterned, brown notodontid moths with a black orbicular spot. Todd (1976) revised the genus whose distribution extends from Mexico to Amazonian Brazil. He described two new species and listed two species, D. angulinea Schaus and D. tuna (Schaus), from Costa Rica. One additional species was recently described by Miller and Thiaucourt (2011) from Ecuador. The genus traditionally has been placed in the Nystaleinae, but that placement remains provisional because species of Dunama lack the characteristic morphological traits of most nystaleines. Additionally, all known caterpillars of Dunama feed on monocots (Musaceae, Marantaceae, Heliconiaceae, Arecaceae), a trait rarely encountered in the Notodontidae. Review of the Costa Rican species is part of an ongoing documentation of over 700 notodontid species collected or reared by parataxonomists and others in Area de Conservación Guanacaste (Janzen 2004, Janzen et al. 2009, and see http://janzen.bio. upenn.edu/caterpillars/database.lasso).

Material and methods
About 1,700 spread specimens of Dunama spp. were examined as follows: 1,545 rearing records from the project "Inventory of the caterpillars of Area de Conservación Guanacaste (ACG), and their parasitoids and food plants" (see Janzen 2004, Janzen et al. 2009 and also search on Dunama spp. at http://janzen. bio.upenn.edu/caterpillars/database.lasso), where the species reside under their interim names until this paper is published. 73 light-caught specimens from the INBio Lepidoptera collection of the project "National Inventory of Biodiversity 1978-2011." 53 light-caught specimens from the collection of J. Bolling Sullivan, Beaufort, NC, USA.
Genital dissections and measurements were made using an Olympus SZ60 stereomicroscope with a calibrated ocular micrometer. The following protocol was used for the dissection of genitalia: abdomens were digested in 10% KOH, cleared, and stained with mercurochrome and Eosin Y (Montero-Ramírez et al. 2011). Genitalia and pelts were stored in glycerol for examination (in 70% ethanol solution, 3:1) and subsequently slide mounted using Euparal. Genital slides were photographed using a JVC 3-CCD color video camera attached to an Olympus SZ60 stereomicroscope, both mounted in an Olympus SZH-ILLD illumination base. Using Montage explorer software (version: 2.01.0075, Synoptics Ltd.) and Auto-Montage software (version: 4.02.0014, Synoptics Ltd.) photographs were enhanced for publication.
Morphological terminology follows Miller (1991). A subset of the total specimens was used for species descriptions.
All holotypes and representative paratypes are deposited in the collections at the Instituto Nacional de Biodiversidad (INBio), Santo Domingo de Heredia, Costa Rica, and the other paratypes are in the USNM.
wing with M1 from proximal third of narrow accessory cell; hindwing with Sc from middle of cell diverging from Rs and straight; Rs and M1 connate from upper angle and M3 and Cu1 connate from lower angle of the cell. Male terminal tergites distinctive. Male genitalia -Uncus short, rounded, sometimes divided; socii sclerotized, upcurved; valves with costal margin sclerotized, sometimes with projections; anal margin often partially but narrowly sclerotized and usually diagnostic; juxta undifferentiated, transtilla membraneous; phallus well developed, sclerotized, usually extending to uncus and narrowing distally; often with distinct lateral and dorsolateral processes; eighth sternum diagnostic, quadrate basally but often with multiple distal projections. Female genitalia -Ovipositor lobes large, often sclerotized, ostia sclerotized, ductus and corpus bursae reduced, membranous and without signa. Larvae -Brightly colored, feeding on monocots. Schaus, 1912 http://species-id.net/wiki/Dunama_angulinea Figs 1-6 Type material. Holotype male: Guapiles, Costa Rica, 17505 USNM (examined).
Diagnosis. Sternum 8 (St8) wide, short, anterior margin simple, posterior margin bearing a pair of small, widely-separated processes. Phallus thin basally, wider medially, with a pair of short, serrate projections on each margin, distal part with a pair of opposite, marginal, non-serrate projections, longer than anterior ones. Vesica short, unsclerotized, without cornuti. The single pair of terminal, widely-separated processes of sternum 8 and the tripartite distal structure of the phallus distinguishes D. angulinea from its known congeners.
Redescription. . Head -Antenna pectinate in basal 4/5, rami moderately long, reddish brown, distal fifth of shaft simple, cream colored with an intermix of reddish-brown and gray-brown scales; scape with scale tuft reddish brown and cream colored; ocelli absent; labial palpus upcurved, reddish brown with a few scattered cream-colored scales; vertex reddish brown, cream colored laterally; patagium reddish brown near midline, reddish brown laterally, margins cream colored. Thorax and abdomen -Tegula cream colored at base, a mix of cream and reddish-brown scales distally; mesoscutum reddish brown anteriorly, cream and reddish brown posteriorly; mesoscutellum mostly creamy white; thoracic pleuron cream colored to reddish brown; legs mostly reddish brown on outer surfaces, cream colored on inner ones. Abdominal dorsum light gray, venter cream colored. Wings -Forewing dorsal ground color a mixture of gray-brown, reddish-brown, and beige-colored scales; veins lined with gray, especially distally; anal fold and cubitus reddish brown; orbicular spot diffuse reddish brown; reniform spot small, reddish brown; medial (M) line thin, reddish brown, a wide, vaguely-defined beige band beyond it; postmedial (PM) line thin, reddish brown, poorly defined; adterminal (AD) line reddish brown, fringe gray brown. Ventral surfaces of both wings gray brown (Figs 1, 2). Dorsal hindwing dirty gray brown, lighter near base. Wing length (WL) 11.20-12.2 mm). Male genitalia (Figs 4-6) -tergum 8 quadrate, posterior margin narrowly sclerotized; St8 wide, short, anterior margin simple, posterior margin extended and bearing a pair of long processes (Fig. 5). Uncus lobule-like, short and pubescent; socci thin, short and slightly curved. Valva broad and membranous, with saccular margin serrate, inner surface with spine-like process, near apex (Fig. 4). Phallus thin basally, wider medially, with a pair of short, serrate projections on each margin, distal part with a pair of opposite marginal non-serrate projections longer than anterior ones. Vesica short, unsclerotized (Fig. 6). Female. Unknown. Natural history. Unknown Distribution. Adults of Dunama angulinea have been collected from Limon Province (Hitoy Cerere, Verugua Rainforest, Guapiles). The distribution follows the Caribbean coast and we know of no records farther inland (Fig. 85). Two specimens identical in size and maculation from La Selva (Heredia) were found on dissection to represent another species described below.
Remarks. Todd (1976, Fig . 10  855 ♂). The holotype from Guapiles, Costa Rica was not dissected. When we dissected the type, it was obvious that the Guatemalan paratypes represent another species. We are not describing the Guatemalan species as new because its genitalia are very similar to those of D. tuna and because we have no specimens from the area between Guatemala and Costa Rica. We also have no barcode or life history data for the Guatemalan species. The barcode NJ tree (Fig. 86) associates Dunama angulinea with D. jessiehillae, described below, from western and inland Costa Rica and can be distinguished from it only by the shape of the sternum. One haplotype represents all three specimens of Dunama angulinea that are recent enough for molecular analysis. Other material examined. Barcoded: 198 specimens that divided into four haplotypes with slight differences from the most common haplotype (163 specimens) of 0.13% or less; we do not consider these differences to be of species-level significance and many are due to slightly shorter barcode sequences. No specimens from Heredia Province were barcoded. Museum specimens: (45 specimens) 2♂ 3♀ Guanacaste, 20♂ 10♀ Alajuela, 10♂ Heredia. Dissections: 1♂ 2♀ Guanacaste, 2♂ 1♀ Alajuela, 2♂ Heredia. INBio, USNM, JBS. Janzen & Hallwachs vouchers of reared specimens.  Etymology. This species is named in honor of Ms. Jessie Hill of Hawaii and Philadelphia, and great-great-grandaughter of Ms. Jessie Barron, and in emphatic recognition of Jessie Hill's contribution to saving and inventorying the conserved ACG rain forest in which reside Dunama jessiehillae and four other new species of Dunama described in this report.

Dunama jessiehillae
Diagnosis. St8 wide, short, anterior margin simple, posterior margin bearing a pair of small, widely separated processes, a second long pair of processes arises between this more basal pair. Phallus thin basally, wider medially, with a pair of short, serrate projections on each margin, distal part with a pair of opposite, marginal, non-serrate projections, longer than anterior ones. Vesica short, unsclerotized, no cornuti. The tripartite distal structure of the phallus and the two pairs of processes on St8 distinguish D. jessiehillae from its known congeners.
Description. Male (Figs 7,8,[11][12][13]. Head -Antenna pectinate in basal 4/5, rami moderately long, reddish brown, distal fifth of shaft simple, cream colored with an intermix of reddish-brown and gray-brown scales; scape with scale tuft reddish brown and cream colored; frons with cream-colored scales interspersed with a few reddish-brown scales, ocelli absent; labial palpus upcurved, reddish brown with a few scattered cream-colored scales; vertex reddish brown, cream colored laterally; patagium reddish brown near the midline, reddish brown laterally, margins cream colored. Thorax and abdomen -Tegula cream colored at base, a mix of cream and reddishbrown scales distally; mesoscutum reddish brown anteriorly, cream and reddish brown posteriorly; mesoscutellum mostly creamy white; thoracic pleuron cream colored to reddish brown; legs mostly reddish brown on outer surfaces, cream colored on inner surfaces. Abdominal dorsum light gray, venter cream colored. Wings -Forewing dor-sal ground color a mixture of gray-brown, reddish-brown and beige-colored scales; veins lined with gray, especially distally; anal fold and cubitus reddish brown; orbicular spot diffuse reddish brown; reniform spot small, reddish brown; medial M line thin, reddish brown, a wide, vaguely-defined beige band beyond it; postmedial PM line thin, reddish brown, poorly defined; adterminal AD line reddish brown, fringe gray brown. Ventral surfaces of both wings gray brown. Dorsal hindwing dirty gray brown, lighter near base (Figs 7,8). (WL 10.70-13.3 mm). Male genitalia (Figs 11-13) -T8 quadrate, posterior margin narrowly sclerotized; St8 wide, short, anterior margin simple, posterior margin bearing a pair of small, widely separated processes, a second, longer pair of processes arise between arms of fork ( Fig. 11). Uncus lobule-like, short and pubescent; socci thin, short and slightly curved. Valva broad and membranous, with saccular margin serrate and inner surface with spine-like process, near appex ( Fig.  12). Phallus thin basally, wider medially with a pair of short, serrate projections on each margin, distal part with a pair of opposite marginal non-serrate projections, longer than anterior ones. Vesica short, unsclerotized ( Fig. 13). Female (Figs 9,10,14). Antenna filiform, shaft cream with a mix of reddish-and gray-brown scales; body color and wing pattern similar to male, wings longer and darker (Figs 9, 10). (WL 13.7-14.7 mm). Female genitalia (Fig. 14) -St8 forming a heavily sclerotized capsule; anterior apophyses thin and acute; posterior apophyses thin, CB small and rounded, signum absent; DB short; ostium recessed in St8. Ovipositor lobes triangulate and setose.
Natural history (Figs 74,79,80,81). 506 rearing records: ACG locations: Sector Del Oro (n=84), Orosi (n=4), Pitilla (n=68), Rincon Rain Forest (n=132), San Cristobal (n=217); all mid-elevation rain forest and does not occur in ACG dry forest either as caterpillars or free-flying adults. Eggs laid in small batches of 5-40, and caterpillars may remain together through the penultimate instar, but generally forage separately in the last instar. Cocoons are solitary, generally in a fold of the palm leaf or two pinnae one on top of the other, lightly silked together. The relatively conspicuous caterpillars remain on the leaf when disturbed rather than drop to the ground, implying that they may be aposematic or mimetic even though they are commonly difficult to encounter among overlapping leaf parts, and often on the underside of the leaf.
Distribution. Adults of Dunama jessiehillae have been collected on the east slope of Cordillera Volcanica de Guanacaste and Tilaran, and in the Sarapiqui lowlands, from 40 to 1500 m elevation ( Fig. 85), but larvae have only been encountered at midelevations on the same slopes.
Remarks. This species feeds exclusively on Arecaceae. Several barcode haplotypes are present in populations from La Selva, Heredia west to the Pacific coast, but they are all very similar (Fig. 86). The most common haplotype is shared with D. angulinea, which ocupies the Caribbean coastal area and differs only in the structure of its sternum. We elected on that basis to describe it as a new species, following both the advice of a reviewer and our own analysis. If later studies support the doubtful hypothesis that this is merely geographic variation in a widely distributed species, then D. jessiehillae would be synonymized with D. angulinea. Other material examined. Barcoded: 25 specimens that divided into 6 apparent haplotypes with differences from the most common haplotype (16 specimens) of 0.3% or less (Fig. 86) Etymology. This species is named in honor of Ms. Jessie Barron, great-greatgrandmother of Jessie Hill of Philadelphia and Hawaii, and in emphatic recognition of Jessie Hill's contribution to saving and inventorying the conserved ACG rain forest in which Dunama jessiebarronae breeds.

Dunama jessiebarronae
Diagnosis. St8 wide, short, anterior margin simple, posterior margin bearing a pair of long, widely separated processes with serrate margins. Phallus thin at the base, wider and sclerotized in the distal part, with a pair of opposite basal projections at the base of the sclerotized portion and a small subterminal projection distally on the left side and a small finger-like terminal projection.
Description. Male (Figs 15,16,[19][20][21]. Head -Antenna pectinate in basal 4/5, rami moderately long, black, distal fifth simple, shaft cream with a few reddish-brown scales, scape with scale tuft reddish brown and creamy; haustellum well developed, frons with a mix of creamy and reddish-brown scales; labial palpus upcurved, blackish brown with a few scattered cream-colored scales; ocelli absent; vertex cream, reddish brown laterally; patagium reddish brown near midline, blackish brown laterally, margins cream colored. Thorax and abdomen -Tegula cream colored at the base, a mix of cream and reddish-brown scales distally, mesoscutum reddish brown anteriorly, cream and reddish brown posteriorly, mesoscutellum mostly creamy white; thoracic pleuron cream colored to reddish brown; legs mostly reddish brown on outer surfaces, cream colored on inner ones. Abdominal dorsum light gray, venter cream colored. Wings -Forewing dorsal ground color cream at base, 2/3 black brown; a prominent blackbrown, slightly oblique bar between base of reniform spot and base of inner margin of forewing in both sexes; veins lined with gray, especially distally; anal fold and cubitus blackish brown; orbicular spot diffuse, blackish brown; reniform spot small, blackish brown; M-line thin, blackish brown, a wide, vaguely-defined beige band beyond it; AD-line with blackish-brown spots, fringe gray brown. Ventral surfaces of both wings gray brown. Forewing costal margin cream ventrally. Hindwing dorsal dirty gray brown, lighter near base (Figs 15, 16). (WL 13.5-13.7 mm). Male genitalia (Figs 19-21) -T8 quadrate, posterior margin narrrowly sclerotized; St8 wide, short, anterior end with simple margin, posterior margin bearing a pair of long, widely-separated processes with serrate margins (Fig. 19). Uncus elongated lobule-like and lightly pubescent, socci thin and curved. Valva broad and membranous, with slightly undulating serrate saccular margin, inner surface with a small spine-like process near apex (Fig.  20). Phallus thin at base, widening and sclerotized distally, with a pair of opposite basal projections at base of sclerotized portion, which has a small subterminal projection on left side and a small finger-like terminal projection (Fig. 21). Vesica small. Female (Figs 17,18,22). Antenna filiform, shaft cream; body color and wing pattern similar to male but wings longer and darker (Figs 17, 18). (WL 16.6-17.2 mm). Female genitalia (Fig. 22)  Parasitoids. Tachinidae: out of 489 rearing records of wild-caught caterpillars, 10 records generated Lespesia Wood33DHJ06, DHJPAR0015889, DHJPAR0015890, DHJPAR0015880, DHJPAR0015878, DHJPAR0015903, DHJPAR0008353, DHJPAR0015879, DHJPAR0015875, DHJPAR0015876, DHJPAR0015882. This fly parasitizes only Notodontidae feeding on Heliconiaceae and Musaceae, and also parasitizes three other species of Dunama feeding on Arecaceae, as well as Dottia and Heorta feeding on the same family. A single record (DHJPAR0016614) of Lespesia Wood03bDHJ05 parasitizing Dunama jessibaronae is not surprising, given that this fly parasitzes five caterpillar families feeding on a large variety of plant species and families, and as such is quite "generalist." Distribution. Dunama jessiebarronae has been collected on the east slope of Cordillera Volcanica de Guanacaste, Cordillera Volcanica Central, Cordillera de Talamanca, llanuras de Sarapiqui, and the lowlands of the Caribbean, from 50 to 1115 m elevation (Fig. 85).
Remarks. Adults of Dunama jessiebarronae have almost the exact maculation pattern of Dunama tuna (Schaus), but the latter moth is larger. In addition, there are two lateral prongs at the base of the posterior projection of the sternum. Todd (1976) made D. sagittula (Dognin, 1914), a Colombian species, a synonym of D. tuna, whose type locality is also in Colombia. Although he did not dissect the type of D. tuna, we did, and it agrees with that of D. sagittula. Todd also reported a male specimen from Porto Bello, Panama and from Sixaola River, Costa Rica. These two Central American specimens are smaller than are those from Colombia, and they also have lateral prongs on the sternum, which are absent in all Costa Rican specimens of D. jessiebarronae we have examined, including 2 males from Hitoy Cerere, Limon Province, which is located not far from Sixaola River. Because we were unable to find any recent specimens with lateral prongs on the tergum, and because barcoding has shown few species in common between Costa Rica and northern South America (unpublished), we have chosen to treat the Costa Rican populations as a new species, D. jessiebarronae. The specimens from Sixaola River and Panama cited by Todd are believed to be D. jessiebarronae. Changes in the form of the sternum along the Atlantic Coast are also seen in the closely related species D. jessiehillae and D. angulinea, albiet it is the reverse change. Future examination and barcoding of specimens collected in Panama and northern Colombia should resolve this issue. Other material examined. Barcoded: 78 specimens, which divided into four haplotypes with differences from the most common haplotype (44 specimens) of 0.25% or less, except for the specimens from Limon that were idéntical, or about 0.6% different. Barcoded specimens were from Alajuela and Limon Provinces (Fig. 86)  Etymology. This species is named in honor of Ms. Jane Waldron, great-grandmother of Jessie Hill of Philadelphia and Hawaii, and in emphatic recognition of Jessie Hill's contribution to saving and inventorying the conserved ACG rain forest in which Dunama janewaldronae breeds.

Dunama janewaldronae
Diagnosis. St8 wide, short, anterior margin simple, posterior margin bearing a pair of small, widely separated processes with serrate margins, a second, long pair of processes arising between arms of first pair. Phallus thin in the base, wider and sclerotized in distal part, ventral margin of distal part with six teeth and dorsal margin with two small teeth.
Description. Male (Figs 23,24,[27][28][29]. Head -Antenna pectinate in basal 4/5, rami moderately long reddish brown, distal fifth simple, shaft with a mix of reddishbrown and gray brown scales, scape with scale tuft blackish brown at base and cream to tip; frons with a mix of cream and blackish-brown scales; labial palpus upcurved blackish brown with a few scattered cream-colored scales; ocelli absent; vertex blackish brown, cream colored laterally; patagium blackish brown near midline, blackish brown laterally, margins cream colored. Thorax and abdomen -Tegula cream colored at base, a mix of cream and blackish-brown scales distally; mesoscutum blackish brown anteriorly, cream and reddish brown posteriorly; mesoscutellum mostly creamy white; thoracic pleuron cream colored to blackish brown; legs mostly blackish brown on outer surfaces, cream colored on inner surfaces. Abdominal dorsum light gray, venter cream colored. Wings -Dorsal ground color a mixture of gray-brown and beige scales; veins lined with gray, especially distally; anal fold and cubitus blackish brown; orbicular spot diffuse blackish brown; reniform spot small, blackish brown; M-line thin, blackish brown, a wide, vaguely-defined beige band beyond it; PM-line thin, blackish brown, poorly defined; AD-line with blackish-brown spots, fringe gray brown. Ventral surfaces of both wings gray brown. Dorsal hindwing dirty gray brown, lighter near base (Figs 23, 24). (WL 12.5-13.4 mm). Male genitalia (Figs 27-29) -St8 wide, short, anterior margin simple, posterior margin bearing a pair of small, widely separated processes with serrate margins; a second, long pair of processes arising between arms of first pair (Fig. 27). Uncus lobule-like and lightly pubescent, socci thin and lightly curved. Valva broad and membranous, with serrate saccular margin, inner surface with a hook-like process near apex (Fig. 28). Phallus thin at base, wider and sclerotized distally, ventral margin of distal part with six teeth and dorsal margin with two small teeth. Vesica tiny, bearing a minute cornutus (Fig. 29). Female (Figs 25, 26, 30). Antenna filiform, shaft gray brown; body color and wing pattern similar to male but wings longer and darker (Figs 25, 26). . Female genitalia (Fig. 30) -Segment 8 forming a heavily sclerotized capsule; anterior apophyses acute; posterior apophyses tiny, CB small and rounded, signum absent; DB short; ostium recessed in St8. Ovipositor lobes triangulate and setose.
Distribution. Dunama janewaldronae has been reared from intermediate elevations of the eastern side of the Cordillera Volcanica de Guanacaste from 400 to 680 m elevation (Fig. 85).
Remarks. This species shows identical genitalia and very similar barcodes throughout Costa Rica (Fig. 86). Specimens from the Caribbean side of Costa Rica have the most divergent barcodes, but are still within the range of variation seen for most species. Nearest neighbor analyses pair D. janewaldronae with D. angulinea and they differ mostly in being slightly different in size. They share several species of understory Arecaceae as caterpillar food plants. Other material examined. Barcoded: 75 specimens from Guanacaste and Puntarenas Provinces that divide into 3 principal haplotypes (excluding 2 partial sequences), which differed from each other by less than 0.25%. One haplotype predominated (66 specimens). Museum specimens: 6 specimens: 3♂ 2♀ Guanacaste, Etymology. This species is named in honor of Ms. Jessie Bancroft, grandmother of Jessie Hill of Philadelphia and Hawaii, and in emphatic recognition of Jessie Hill's contribution to saving and inventorying the conserved ACG rain forest in which Dunama jessiebancroftae breeds.

Dunama jessiebancroftae
Diagnosis. St8 wide and short, anterior margin simple, posterior margin bearing a simple acute and triangulate process. Phallus narrow at base, expanding medially, heavily sclerotized at distal third, with dorsal margin serrate, narrowing to tip.
Description. Male (Figs 31,32,[35][36][37]. Head -Antenna pectinate in basal 4/5, rami moderately long reddish brown, distal fifth simple, shaft gray brown with reddish-brown scales at base, scape with scale tuft gray brown and cream; frons with a mix cream and gray-brown scales; labial palpus upcurved blackish brown with a few scattered cream-colored scales; ocelli absent; vertex gray brown, cream colored laterally; patagium blackish brown with margins cream colored. Thorax and abdomen -Tegula cream colored at base, a mix of cream and gray-brown scales distally; mesoscutum blackish brown anteriorly, cream and blackish brown posteriorly; mesoscutellum mostly creamy white; thoracic pleuron cream colored to blackish brown; legs mostly reddish brown on outer surfaces, cream colored on inner surfaces. Abdominal dorsum light gray, venter cream colored. Wings -Dorsal ground color with a mixture of graybrown, blackish-brown and beige scales; veins lined with gray, especially distally; anal fold and cubitus blackish brown; orbicular spot diffuse blackish brown; reniform spot small, blackish brown; M-line thin, wavy, blackish brown, a wide, vaguely-defined beige band beyond it; PM-line thin, blackish brown, poorly defined; AD-terminal line with blackish-brown spots, fringe gray brown. Ventral surfaces of both wings gray brown. Dorsal hindwing dirty gray brown, lighter near base (Figs 31, 32). Male genitalia - (Figs 35-37) St8 wide and short, anterior margin simple, posterior margin bearing a simple acute and triangulate process (Fig. 35). Uncus small with a hollow depression in middle, socci thin, long and acute. Valva wide and membranous with saccular margin serrate, heavily sclerotized at base (Fig. 36). Phallus narrow at base, expanding medially, heavily sclerotized at distal third, with dorsal margin serrate, narrowing to tip. Vesica tiny (Fig. 37). Female (Figs 33,34,38). Antenna filiform, shaft gray brown with a mix of reddish-brown scales; Body color and wing pattern similar to male but wings longer and darker (Figs 33, 34). .0 mm). Female genitalia (Fig. 38) -Segment 8 forming a heavily sclerotized capsule; anterior apophyses thin and acute; posterior apophyses thin, CB small and round, signum absent; DB short; ostium recessed in St8. Ovipositor lobes acute and setose.

Parasitoids. Braconidae:
Macrocentrinae? Austrozele? (n=1); Meteorinae, Meteorus Zitani01DHJ05 (n=1). Tachinidae: Lespesia Wood33DHJ06 (n=6), a species of tachinid parasitoid that it shares with two other species of Dunama. Distribution and habitat. In addition to the rearing records from ACG, Dunama jessiebancroftae has been collected in the Peninsula de Nicoya, and the lowland of Central Pacific Costa Rica, from 50 to 1286 m elevation (Fig. 85); all of these extra-ACG sites are also intergrades between rain forest and dry forest, at least before they were largely deforested.
Remarks. This species is homogeneous over its limited range. Nearest neighbor analyses (Fig. 86) suggest that it is the most different from all other Dunama in Costa Rica, which suggests the highly unlikely scenario that it was orginally a species of the intergrade of dry forest with rain forest, and then evolutionarily spread into rain forest ecosystems. Other material examined: Barcoded: 22 specimens that segregated into four haplotypes with differences from the most common haplotype from Alajuela (14 specimens) of 0.1% for a single haplotype from Alajuela, 1.1% for four specimens from Puntarenas, and 1.4% for three specimens from Cartago. The Cartago and Puntarenas specimens differed by 0.6%. Musem specimens: (14 specimens) 2♂ 1♀ Alajuela, 8♂ Cartago, 2♂ Puntarenas. Dissections: 1♂ Alajuela, 2♂ l♀ Guanacaste, 3♂ l♀ Cartago, 2♂ Puntarenas. Etymology. This species is named in honor of Ms. Jane Cox, mother of Jessie Hill of Philadelphia and Hawaii, and in emphatic recognition of Jessie Hill's contribution to saving and inventorying the conserved ACG rain forest in which Dunama janecoxae breeds.

Dunama janecoxae
Diagnosis. St8 wide, short, anterior margin simple, posterior margin densely sclerotized with a rectangular shape lacking any processes; phallus simple, thin at base, heavily sclerotized distally part, with a pair of small triangular projections subopposite on each margin. The medial projection from the costa of the male genitalia is unique among species of Dunama.
Distribution. Dunama janecoxae is the upper elevational species of Dunama on the Cordillera Volcanica de Guanacaste, and the eastern slope of Cordillera de Tilaran and Talamanca, occurring from 1090 to 1185 m elevation (Fig. 85).
Remarks. Dunama janecoxae seems to offer the classic conundrum of isolated populations in the upper elevations of isolated mountains. Each population has a unique and slightly different barcode (Fig. 86), but the genitalia differ only slightly among populations and far less than that displayed among most other species of Dunama in Costa Rica. In as much as we have life history data for only the ACG population, we elect to leave these mountaintop populations as one species, even though their morphological and barcode differences are of the same degree as other ACG sympatric/parapatric pairs of species with distinct but similar barcodes (e.g., Janzen et al. 2005; Neoxeniades luda (Hewitson) and N. pluviasilva Burns (Burns et al. 2007); four sympatric/parapatric species of Perichares Scudder (Burns et al. 2008)). Additional material will be needed to determine the extent of separation of these different montane populations. Etymology. Dunama biosise is named in honor of BIOSIS, the non-profit publishing company, the sale of which generated the JRS Biodiversity Foundation (http:// www.jrsbdf.org), which in turn supports biodiversity information management for conservation in many places, including INBio and ACG.

Dunama biosise
Diagnosis. St8 wide, short, anterior margin simple, posterior margin sclerotized with a pair of forceps-like processes, a small and sclerotized triangular projection at the base of each process. Phallus with subbasal, unsclerotized expansion, distal half narrow and sclerotized.
Natural history. Unknown, except that it is a moth of Costa Rica's lowland Pacific coast rain forest and both sexes can be captured at light at night.
Distribution. Dunama biosise has been collected from 0 to 100 m elevation in the Osa Peninsula, Area de Conservacion Osa (Fig. 85).
Distribution and habitat. Dunama indereci has been collected only in Villa Blanca, in San Ramon, Alajuela province, at 1115 m elevation, in a montane pass between Costa Rica's Cordillera de Tilaran and Volcanica Central (Fig. 85).
Remarks. This species feeds exclusively on Heliconia latispatha. One barcode haplotype was recored in the population from Villa Blanca (Fig. 86).  The Dunama species from Costa Rica in an NJ barcoding tree. Sample sizes are restricted to a haphazardly selected set of five specimens for each species. Many specimens were reared from wildcaught caterpillars and further information on each can be found at Janzen and Hallwachs (2012).