Nine new species of Uramya Robineau-Desvoidy (Diptera: Tachinidae) from Area de Conservación Guanacaste in northwestern Costa Rica, with a key to their identification

Abstract Background We describe nine new species in the genus Uramya Robineau-Desvoidy, 1830 from Area de Conservación Guanacaste (ACG) in northwestern Costa Rica. All species were reared from an ongoing inventory of wild-caught caterpillars spanning a variety of families (Lepidoptera: Erebidae; Limacodidae; Megalopygidae; Lasiocampidae and Dalceridae). Our study provides a concise description of each new species using morphology, life history, molecular data, and photographic documentation. In addition to the new species the authors provide a redescription the previously described Uramya sibinivora Guimarães, which was also collected within ACG during this study. We also provide a redescription of the genus, and a revised key to species of Uramya occurring in Central and South America. New information The following nine new species of Uramya, all authored by Fleming & Wood, are described: Uramya albosetulosa Fleming & Wood sp. nov., Uramya constricta Fleming & Wood sp. nov., Uramya contraria Fleming & Wood sp. nov., Uramya infracta Fleming & Wood sp. nov., Uramya lativittata Fleming & Wood sp. nov., Uramya lunula Fleming & Wood sp. nov., Uramya nitida Fleming & Wood sp. nov., Uramya pannosa Fleming & Wood sp. nov., and Uramya penicillata Fleming & Wood sp. nov. The following are proposed by Wood as new synonyms of Uramya: Olinda Townsend, syn. nov. and Procleonice Townsend, syn. nov. The following new combination is proposed as a result of the new synonymies: Uramya brasiliensis Macquart, comb. nov. Procleonice prolixa Townsend is synonymized under Uramya brevicauda Curran, syn. nov.


Introduction
Tachinidae are the second most diverse family of Diptera (Belshaw 1993, O'Hara 2008, with almost 10,000 described species classified into over 1500 genera (O'Hara 2008, O'Hara 2014. The number of named species catalogued by Guimaraes (1971) for the Neotropical Region, some 2,864 species, is larger than that of any other region. Based on what has recently been discovered in Costa Rica and what is already present in other collections, this number is undoubtedly just a small fraction of what actually exists in nature. The most speciose genera of the Costa Rican tachinid fauna seem to occur in the upper elevations and cloud forests that extend from the western slopes of the Sierra Madre Occidental in Mexico to both slopes of the Andes, from Colombia south to Bolivia. The present study describes 9 new Neotropical species of Uramya Robineau-Desvoidy, 1830 (Dexiinae: Uramyini) from Area de Conservación Guanacaste (ACG) in northwestern Costa Rica (http://www.acguanacaste.ac.cr) and provides a key to their identification and that of their Central and South American congeners.
The last major taxonomic work on the Uramyini was by : his "Revision of the South American Uramyini (Diptera, Tachinidae)" provided a concise and complete diagnosis of the tribe and included two nominal genera: Uramya and Thelairaporia . Robineau-Desvoidy (1830 erected the genus Uramya based on one male collected in Brasil, misidentified as a female, which he named U. producta Robineau-Desvoidy, 1830. Like other genera within the Uramyiini (e.g., Itaplectops Townsend, 1927 and Thelairaporia), species of Uramya parasitize caterpillars within the Arctiinae (Erebidae), Limacodidae, Megalopygidae, Lasiocampidae, and Dalceridae (Arnaud 1978, Wood and Zumbado 2010. All flies and rearing information described here derive from the ongoing inventory of the tritrophic relationships between caterpillars, their food plants and their parasitoids within the dry, rain, and cloud forests of the terrestrial portion of ACG , Janzen and Hallwachs 2011, Rodriguez et al. 2012, Fleming et al. 2014b, Janzen and Hallwachs 2015. Since 1978 this inventory has yielded an unprecedented amount of invaluable information on the tri-trophic relationships between parasitoids, hosts and associated food plants , Janzen and Hallwachs 2011, Janzen and Hallwachs 2015, Fernandez-Triana et al. 2014).
Our descriptions of these nine new species of Uramya build on existing knowledge and are based on differences in external morphology, COI (coxI or cytochrome c oxidase I) gene sequences, and male terminalia (when necessary). As the inventory is continually growing, it should be noted that this paper should not be taken as an indication of the final total number of species of Uramya present in ACG or Costa Rica. Our descriptions are limited only to the species known and reared from ACG. This paper on Uramya is part of a larger effort to describe new species reared during the ACG inventory (Fleming et al. 2014a, Fleming et al. 2014b, Fleming et al. 2015c, Fleming et al. 2015d, Fleming et al. 2016a, Fleming et al. 2016b). This series of taxonomic papers will represent a baseline for later, detailed ecological and behavioral accounts and studies extending across ACG ecological groups, whole ecosystems, and taxonomic assemblages much larger than a genus.

Project aims and rearing intensity
All reared specimens were obtained from host caterpillars collected in ACG , Janzen and Hallwachs 2011, Janzen and Hallwachs 2015. ACG's 125,000+ terrestrial hectares span the provinces of Alajuela and Guanacaste, along the dry forested northwestern coast of Costa Rica and inland to the Caribbean lowland rain forest. ACG comprises several different biomes and intergrades, ranging from sea level up to 2,000 m. The tachinid rearing methods are described at http://janzen.bio.upenn.edu/caterpillars/ methodology/how/parasitoid_husbandry.htm. Since its inception, this inventory has reared over 750,000 wild-caught ACG caterpillars. Any frequencies of parasitization reported here need to be considered against this background inventory. Comparative details of the parasitization ecology of these flies will be treated separately in later papers, in the context of the study of all parasitization rates of tachinids on ACG caterpillars, once the overall alpha taxonomy of ACG caterpillar-attacking tachinids is more complete than at present.

Descriptions and imaging
Species accounts presented in this paper are deliberately brief and only include basic descriptions of body morphology and coloration commonly used in the identification of Tachinidae. The descriptions are complemented with a series of color photos of every species, used to illustrate the morphological differences among them. The morphological terminology used follows Cumming and Wood (2009). All dissections and photography were carried out following the methods detailed in Fleming et al. (2014a). Measurements and examples of parts of the terminalia are illustrated in Fig. 1.

Voucher specimen management
All caterpillars reared from the ACG efforts receive a unique voucher code in the format yy-SRNP-xxxxx. Any parasitoid emerging from a caterpillar receives the same voucher code as a record of the rearing event. If and when the parasitoid is later dealt with individually it receives a second voucher code unique to it, in the format DHJPARxxxxxxx. These voucher codes assigned to both host and parasitoids may be used to obtain the individual rearing record at http://janzen.bio.upenn.edu/caterpillars/database.lasso.

Description
Head: frontal bristles not extending beyond level of pedicel; eyes haired; facial margin not visible in profile; vibrissa arising at level of facial margin; first flagellomere almost reaching facial margin; arista slightly pubescent; facial ridge devoid of bristles; ocellar bristles weak to absent; inner vertical bristles elongate, parallel, in some cases indistiguishable from upper frontal bristles; outer orbitals weak to absent in males, but strong in females; females with two pairs of proclinate orbital bristles.
Abdomen: narrowed and elongate, at least twice as long as wide; frequently acutely produced dorsally into a tail-like process; mid-dorsal depression on ST1+2 reaching hind margin of syntergite; T3, T4 and sometimes ST1+2 with one to three pairs of discal bristles; all tergites with marginals. In females, abdomen usually not as elongate as males and with a rounded posterior end.

Distribution
Nearctic and Neotropical (not known from Chile and southern Argentina)

Ecology
According to Arnaud (1978)Uramya species parasitize lepidopteran larvae in the families Limacodidae, Arctiidae, Megalopygidae and Lasiocampidae. Current data from ACG inventoried larvae confirm this, adding records from the family Dalceridae. Uramya puparia are asymmetrical as seen in Fig. 2 (Fig. 3). Length: 9-14 mm. Head (Fig. 3b): antenna: medial surface of first flagellomere dark orange along upper margin, closest to pedicel; pedicel black; arista 1.5X as long as first flagellomere, dark brown and minutely pubescent; palpus dark yellow, haired; fronto-orbital plate, parafacial and gena silver pollinose; gena bearing few fine hairs along lower margin. Thorax (Fig. 3a, c): entirely silver pollinose; dorsum of thorax and scutellum with conspicuous yellow-white hairs covering surface; sternopleura, hypopleura, pteropleura and ventral surface of abdomen yellow-white pilose; 2 katepisternal bristles; 3 postsutural supra-alar bristles; postpronotum bearing fine black hairs and anepisternum with fine yellow-white hairs; scutellum bearing 1 pair of discal bristles; underside of scutellum bearing a tuft of white hairs near basal marginal bristle. Legs: reddish-yellow in ground color; femur covered in long yellow hairs interspersed among darker hairs and bristles; tarsus all black. Wings: smoky gray translucent; wing veins not infuscate. Abdomen (Fig. 3a): 1 pair of median marginal bristles on ST1+2 and T3; row of marginal bristles on T4 and T5; median discal bristles on T3, T4 and T5; abdomen brown-orange dorsally, with silver pollen on anterior half of T3, T4 and T5; underside of abdomen pale pilose and entirely covered in silver pollinosity. Terminalia (Fig. 3d, e, f): sternite 5 consisting of two small lobes; inner margin covered in dense pollinosity appearing darker than surrounding cuticle; apical edges of lobes of sternite 5 bearing many long, stout, outwardly pointing bristles interspersed with longer bristles close to anterior margin; sternite 5 with wide V-shaped median cleft 0.34X the length of the sternite from lobe apex to base; cercus sharply pointed and distinctly tapered; apical section subequal in length to upper lobe of cercus; slightly curved when viewed laterally, with a slight upward hook at its tip; surstylus oblong, curved and scythe-like in lateral view; posterodorsal half haired, with few short apical bristles; tip of surstylus not lobed when viewed dorsally; in dorsal view, surstylus angled inwards and 1.3X as long as cercus.
Female (Fig. 4). Length: 7-10 mm. As male, except wing smoky translucent amber color and wing veins slightly infuscate; ground color of abdomen as in male except T5 all black and tergal margins appearing yellow on ventral surface.

Diagnosis
Uramya albosetulosa can be distinguished from all other Neotropical species of Uramya by the following combination of traits: dark brown to black antennae, 3 postsutural supra-alar bristles, underside of scutellum with a tuft of white hairs near basal marginal bristle, 1 pair of median marginal bristles on ST1+2 and T3; T4 and T5 with a row of marginal bristles; underside of abdomen pale pilose and silver pollinose.

a b
c d e f

Etymology
The specific epithet is derived from the Latin adjective "albus", for white, and the noun " seta", for bristle, in reference to the tuft of white hairs present along the underside of the scutellum in this species.

Ecology
Uramya albosetulosa sp. nov. has been reared 35 times from Acharia sarans (Dyar), Acharia hyperoche Dognin and Acharia apicalis (Dyar) (Limacodidae), from 2,035 wildcaught mixed siblings and non-siblings found in both dry and rain forest.  (Fig. 5a, c): entirely gray pollinose; dorsum of thorax and scutellum black pilose; sternopleura, hypopleura, pteropleura, and ventral surface of abdomen white pilose; 2 katepisternal bristles; 3 postsutural supra-alar bristles; postpronotum and anepisternum bearing fine black hairs; underside of scutellum bearing a tuft of black hairs near basal marginal bristle. Legs: fore femur gray pollinose, densely covered in thin dark hairs; mid and hind femora dark brown to black in ground color, contrasting with the dark orange tibiae. Wing: smoky grey transparent; wing veins not infuscate. Abdomen (Fig. 5a): median marginal bristles present on all abdominal tergites (ST1+2, T3, T4 and T5); 1 pair of median discal bristles on T3, T4 and T5; ground color of abdomen brown-black up to tergite 5; silver pollinosity present on anterior half of T3, T4 and T5; ST1+2 with lateral gray pollinose spots on posterior half; base of ST1+2 slightly constricted, giving the abdomen a basaly tapered appearance. Terminalia (Fig. 5d, e, f): sternite 5 with two small lobes, inner margin covered in dense pollinosity, appearing darker than surrounding cuticle; apical edges of lobes of sternite 5 bearing short, stout bristles interspersed with longer bristles close to apical margin; sternite 5 with wide Vshaped median cleft, 0.4X the length of sternite from lobe apex to base; cercus sharply pointed, distinctly tapered; apical section 1.4X as long as upper lobe; strongly curved when viewed laterally and with a slight upward hook at its tip; surstylus equilaterally oblong and scythe-like in lateral view; posterodorsal half densely haired; apex of surstylus with a slight lobe when viewed dorsally and few short apical bristles; surstyli weakly angled inwards in dorsal view, almost parallel; surstylus 1.6X as long as cercus.

Diagnosis
Uramya constricta can be distinguished from all other Neotropical species of Uramya by the following combination of traits: pedicel dark brown, 3 postsutural supra-alar bristles, underside of scutellum with a tuft of black hairs near basal marginal bristle, only 1 pair of median marginal bristles on ST1+2, T3 and T4, and by the shape of the surstylus.

Etymology
The specific epithet is derived from the Latin adjective "constrictus", for constricted or compressed, in reference to the slightly constricted base of ST1+2 where it meets the thorax.

Ecology
Uramya constricta has been reared 38 times from Acharia ophelians Dyar (Limacodidae) (one sibling brood of caterpillars) and once from Euclea mesoamericana DHJ04 (Lepidoptera: Limacodidae) in ACG cloud forest. a b Figure 6. Description Male (Fig. 7). Length: 12 -13 mm. Head (Fig. 7b): antenna black, with orange medially and at base, adjacent to pedicel; pedicel black; arista 1.5X as long as first flagellomere, dark brown and minutely pubescent; palpus dark yellow and haired; fronto-orbital plate, parafacial and gena silver pollinose; gena bearing fine hairs along lower margin. Thorax (Fig. 7b, c): entirely gray pollinose; dorsum of thorax and scutellum covered by conspicuous yellow-white hairs; sternopleura, hypopleura, pteropleura, and ventral surface of abdomen yellow-white pilose; 2 katepisternal bristles; 3 postsutural supraalar bristles, 2nd postsutural supra-alar 4X as long as first; postpronotum bearing fine black hairs and anepisternum with fine yellow-white hairs; scutellum bearing 1 pair of discal bristles; underside of scutellum bearing a tuft of white hairs near basal marginal bristle. Legs: reddish-yellow in ground color; femora covered in long, yellow hairs interspersed among darker hairs and bristles; tarsi all black. Wing: smoky gray translucent; wing veins strongly infuscate; infuscation of wing veins becoming more generalized and blending together to a dark gray tone along basal portion of the wing. Abdomen (Fig. 7a): 1 pair of median marginal bristles on ST1+2 and T3; row of marginal bristles on T4 and T5; 2 pairs of median discal bristles on T3 and 1 pair on T4; abdomen dark brown dorsally, with silver pollen on anterior half of T3, T4 and T5; ST1+2 with 2 pollinose spots on either side of mid-dorsal depression; underside of abdomen entirely covered in silver pollinosity. Terminalia (Fig. 7d, e, f): sternite 5 with two small lobes; inner margin covered in dense pollinosity, appearing darker than surrounding cuticle; apical edges of lobes of sternite 5 bearing many long, stout, outwardly pointed bristles interspersed with longer bristles close to lobe margin; sternite 5 with wide V-shaped median cleft, 0.32X length of sternite from lobe apex to base; cercus sharply pointed and distinctly tapered; apical section 1.7X length of upper lobe; slightly curved when viewed laterally, with a slight upward hook at its tip; surstylus narrow, curved and scythe-like in lateral view; middle third of surstylus haired, otherwise almost bare; tip of surstylus not lobed when viewed dorsally and distinctly inwardly angled; surstylus 0.9X as long as cercus.  (Fig. 8). Length: 8-11 mm. As male, except with a slight golden tinge around the thoracic suture, one pair of median discal bristles on T3, T4 and T5, wing smoky brown translucent, only slightly infuscate around major wing veins.

Diagnosis
Uramya contraria can be distinguished from all other Neotropical species of Uramya by the following combination of traits: dark brown to black antennae, 3 postsutural supraalar bristles, 2 strong lateral scutellar bristles, no discal scutellar bristles, underside of scutellum with a tuft of white hairs near basal marginal bristle, abdomen flattened dorsoventrally, ST1+2 with lateral white pollinose spots on either side of mid-dorsal depression, T5 subtriangular, not strongly produced into a long, tail-like process, 1 pair of median discal bristles on T3, T4 and T5, and silver pollinosity on underside of abdomen.

Etymology
The species epithet is derived from the latin adjective " contrarius " for contrary or opposed, referring to its overall similarity but differing terminalia to U. halisidotae.

Ecology
Uramya contraria has been reared four times from a sample of 100 wild-caught, nonsibling Caviria regina Cramer (Erebidae, Lymantriinae) in ACG rain forest.  Description Male (Fig. 9). Length: 14 mm. Head (Fig. 9b): antenna: medial surface of first flagellomere with a dark orange tinge along upper margin, closest to pedicel; pedicel black to dark brown; arista 1.5X as long as first flagellomere, dark brown and minutely pubescent; palpus dark yellow and haired; fronto-orbital plate, parafacial and gena silver pollinose; gena with few fine hairs along lower margin; frontogenal suture darkened but not black. Thorax (Fig. 9a, c): entirely gray pollinose; dorsum of thorax and scutellum with conspicuous black hairs covering surface; sternopleura, hypopleura, pteropleura and ventral surface of abdomen yellow-white pilose; 3 katepisternal bristles; 3 postsutural supra-alar bristles, 2nd postsutural supra-alar 3X as long as first; postpronotum and anepisternum bearing fine black hairs; scutellum bearing 1 pair of almost indistinct discal bristles, and 3 strong lateral marginal bristles; underside of scutellum bearing a tuft of black hairs near basal marginal bristle. Legs: black in ground color, femora covered in long yellow hairs interspersed around femoral bristles; tibiae of dark yellow ground color; tarsi all black. Wing: smoky clear translucent; wing veins slightly infuscate. Abdomen (Fig. 9a): 1 pair of median marginal bristles on ST1+2; row of marginal bristles on T3 and T4; median discal bristles on T3 and T4; black ground color, with silver pollen on either side of mid-dorsal depression on ST1+2; silver pollen on anterior half of T3 and T4; T5 with 2 silver pollinose spots on either side; silver pollinosity extending to underside of abdomen on T3 and T4. Terminalia (Fig. 9d, e, f): sternite 5 with two small lobes, inner margin covered in dense pollinosity appearing slightly darker than surrounding cuticle; apical edges of lobes of sternite 5 bearing many long, stout, outwardly pointed bristles interspersed among shorter hairs, with longer bristles closest to lobe margin; sternite 5 with wide Y-shaped median cleft, 0.48X length of sternite from lobe apex to base; cercus sharply pointed, distinctly tapered; apical section 1.6X length of upper lobe; strongly curved downwards when viewed laterally, lacking any upward hook at its tip; surstylus narrow, curved downwards, and scythe-like in lateral view; surstylus haired along almost its entire length, with tip not lobed when viewed dorsally; in dorsal view, surstyli angled medially so that tips are almost pointing inwards; surstylus 1.3X as long as cercus.

Diagnosis
Uramya infracta can be distinguished from all other Neotropical species of Uramya by the following combination of traits: dark brown to black antennae with only a slight orange tinge and dark brown pedicel, thorax with black hairs interspersed among the bristles dorsally, and yellow-white hairs on sides and ventrally, underside of scutellum with a tuft of black hairs near basal marginal bristle, 1 pair of median marginal bristles on ST1+2, and a row of marginal bristles on T3 and T4.

Etymology
The species epithet is derived from the latin adjective " infractus " meaning broken or weakened, in reference to the broken band of silver pollinosity on T5, which appears as two pollinose spots.

Ecology
Uramya infracta has been reared only once, from a total of 153 wild-caught non-sibling Natada fusca Druce (Limacodidae) caterpillars in ACG rain forest.  Description Male (Fig. 10): 10-15 mm. Head (Fig. 10b): antenna, first flagellomere all black, except orange along upper margin, adjacent to pedicel; pedicel light brown; arista 1.5X as long as first flagellomere, dark brown and minutely pubescent; palpus yellow and haired; fronto-orbital plate, parafacial and gena silver pollinose; gena bearing few fine hairs along lower margin. Thorax (Fig. 10a, c): entirely gray pollinose; surfaces of dorsum of thorax and scutellum with conspicuous black hairs; sternopleura, hypopleura, pteropleura, and ventral surface of abdomen black pilose; 3 katepisternal bristles; 3 postsutural supra-alar bristles, 2nd postsutural supra-alar bristle 4X as long as 1st; postpronotum bearing fine black hairs and anepisternum with fine, yellow-white hairs; scutellum bearing 1 pair of nearly indistinct discal bristles, slightly thicker than scutellar hairs; anatergite bearing a small patch of black hairs; with 2-3 pairs of strong marginal bristles in addition to basal and subapical bristles; underside of scutellum bearing a tuft of black hairs near basal marginal bristle. Legs: femora reddish-brown ground color; tibiae yellow; coxae of a reddish color; tarsi all black. Wing: dark smoky brown translucent; major wing veins strongly infuscate. Abdomen (Fig. 10a): 1 pair of median marginal bristles on ST1+2, T3, T4 and T5; 2 pairs of median discal bristles on T3, T4 and T5; ground color of abdomen black, with silver pollen on basal half of T3, T4, and T5; dorsal surface of tergites not covered by silver pollen, rather bearing a brown pollinosity visible under the microscope, appearing glabrous to the naked eye; no pollen of any sort on ST1+2; underside of abdomen entirely covered in silver pollen, and black hairs. Terminalia (Fig. 10d, e, f): sternite 5 with two broad rectangular lobes; inner edges of lobes of sternite 5 of lobes covered in dense pollinosity, and haired marginally; sternite 5 with a narrow V-shaped median cleft, 0.43X length of sternite from lobe apex to base; cercus long, thin and sharply pointed; slightly curved when viewed laterally, with a slight upward hook at its tip; distinctly tapered, with apical section 1.6X as long as upper lobe; surstylus long and digitiform in lateral view, haired along its entire length, these hairs becoming thicker basally; tip of surstylus not lobed when viewed dorsally; surstylus inwardly curved, 1.1X as long as cercus.

Diagnosis
Uramya lativittata can be distinguished from all other Neotropical species of Uramya by the following combination of traits: dark brown to black antennae, 3 postsutural supraalar bristles, underside of scutellum with a tuft of black hairs near basal bristle, 4-5 pairs of marginal scutellar bristles, and silver pollinosity on underside of abdomen.

Etymology
The species epithet is derived from a combination of the latin nouns " latus ", meaning side, and " vitta ", meaning band, in reference to its distinctive abdominal pattern.
a: habitus in dorsal view b: habitus in lateral view postsutural supra-alar bristles, 2nd postsutural supra-alar 4X as long as 1st postsutural supra-alar; postpronotum and anepisternum bearing fine black hairs; scutellum bearing one pair of discal bristles; underside of scutellum bearing a tuft of black hairs near basal marginal bristle. Legs: entirely black; femora silver pollinose anterodorsally, covered in long black hairs interspersed among darker hairs and bristles. Wings: smoky gray translucent, brownish infuscate costobasally; wing veins not strongly infuscate; infuscation only visible on anterior half of R and M. Abdomen (Fig. 12a): 1 pair of median marginal bristles on ST1+2 and T3, row of marginal bristles on T4 and T5; 2 pairs of median discal bristles on T3, T4 and T5; ground color of abdomen black; posterior margin of ST1+2 silver pollinose; silver pollinose crescent-shaped markings midway along length of tergites T3 and T4; T5 bearing 2 silver pollinose blotches; underside of ST1+2, T3, and T4 silver pollinose; underside of T5 darker. Terminalia: not dissected, as we did not want to dissect the holotype and only known specimen of this species.

Diagnosis
Uramya lunula can be distinguished from all other Neotropical species of Uramya by the following combination of traits: dark brown to black antennae, 3 postsutural supra-alar bristles, 2 strong lateral scutellar bristles, weak discal scutellar bristles, underside of scutellum with a tuft of black hairs near basal marginal bristle, abdomen flattened dorsoventrally, ST1+2 lacking silver pollinose spots on either side of mid-dorsal depression, T5 subtriangular, not strongly produced into a long, tail-like process, 2 pairs of median discal bristles on T3, T4 and T5, and silver pollinosity on underside of abdomen.

Etymology
The species epithet is derived from the Latin noun "lunula", for little moon, in reference to the crescent-shaped markings present on the abdomen of this species.

Ecology
Uramya lunula has been reared only once, from a Podalia orsilocha (Cramer) (Megalopygidae) caterpillar found in ACG rain forest. Podalia orsilocha has been reared 175 times from sibling and non-sibling groups in ACG. apical margin; sternite 5 with wide V-shaped median cleft, 0.6X length of sternite from lobe apex to base; cercus sharply pointed, distinctly tapered; apical section of cercus 2.4X as long as upper lobe; not strongly curved when viewed laterally, with only a slight upward hook at its tip; surstylus oblong, curved and scythe-like in lateral view, with short bristles covering almost its entire surface, and with tips not lobed when viewed dorsally; surstyli not angled inwards, parallel; surstylus 1.2X as long as cercus.

Diagnosis
Uramya nitida can be distinguished from all other Neotropical species of Uramya by the following combination of traits: light-colored, orange-brown antennae, 2 postsutural supra-alar bristles, underside of scutellum with a tuft of black hairs near basal marginal bristle, 1 pair of median marginal bristles on ST1+2, and a row of marginal bristles on T3 and T4.

Etymology
Uramya nitida is derived from the Latin noun "nitidus", meaning bright or glossy, in reference to the glossy appearance of the abdomen under certain angles of light.

Ecology
Uramya nitida has been reared 18 times from Parasa sandrae Corrales & Epstein (Limacodidae), in a sample of 497 non-sibling, wild-caught larvae in both cloud and rain forest.  Description Male (Fig. 15): 15 mm. Head (Fig. 15b): antenna: first flagellomere dark brown on lower 2/3,turning to brownish orange on upper third and adjacent to pedicel; pedicel dark orange; arista 1.5X as long as first flagellomere, dark brown and minutely pubescent; palpus dark orange and haired; fronto-orbital plate, parafacial and gena silver pollinose; gena silver pollinose with a cluster of hairs along lower 1/3 to 1/2; facial ridge darkened, with few fine hairs along lower margin; frontogenal suture black. Thorax (Fig. 15a, c): entirely gray pollinose; dorsum of thorax and scutellum black pilose; sternopleura, hypopleura, pteropleura, and ventral surface of abdomen yellowwhite pilose; 2 katepisternal bristles; 2 postsutural supra-alar bristles; postpronotum and anepisternum bearing fine black hairs; underside of scutellum bearing a tuft of black hairs near basal marginal bristle. Legs: fore femur gray pollinose, densely covered in thin dark hairs interspersed among the bristles; mid and hind tibiae dark brown to black basally, apically dark orange. Wings: smoky gray, with dense infuscation around each vein. Abdomen (Fig. 15a): median marginal bristles only on ST1+2; 2 pairs of median discal bristles on T3 and T4; ground color of abdomen brown-black up to tergite 5; silver pollinosity present on anteriormost half of T3, T4 and T5; ST1+2 with lateral gray pollinose spots on posterior half; band of short black hairs surrounding discal bristles on T3 and T4, abdomen otherwise appearing devoid of hairs. Terminalia (Fig. 15d, e, f): sternite 5 with two small lobes; inner margin covered in dense pollinosity, appearing darker than surrounding cuticle; apical edges of lobes of sternite 5 bearing many medium-length, stout bristles interspersed with longer bristles close to apical margin; sternite 5 with wide V-shaped median cleft, 0.42X length of sternite from lobe apex to base; cercus sharply pointed, distinctly tapered; apical section of cercus 1.4X as long as upper lobe; strongly curved when viewed laterally with a slight upward hook at its tip; surstylus oblong, curved and scythe-like in lateral view; posterodorsal half of surstylus haired, apex with almost no short apical bristles; tip of surstylus not lobed when viewed dorsally; surstylus strongly angled inwards, 1.4X as long as cercus.

Diagnosis
Uramya pannosa can be distinguished from all other Neotropical species of Uramya by the following combination of traits: pedicel orange, 2 postsutural supra-alar bristles, underside of scutellum with a tuft of black hairs near basal marginal bristle, overall absence of abdominal hairs, only 1 pair of median marginal bristles on ST1+2, and 2 pairs of median discal bristles on T3 and T4.

Etymology
The species epithet is derived from the Latin adjective " pannosus ", meaining tattered or shabby, in reference to the irregular and ragged appearance of the abdominal banding.

Ecology
Uramya pannosa has been reared 5 times from Parasa macrodonta Hering & Hopp (Limacodidae) in both cloud and rain forest, and is the only species of tachinid to have been reared from this species of caterpillar in 166 non-sibling rearings. a b Figure 16. Description Male (Fig. 17): 10 mm. Head (Fig. 17b): antenna: pedicel orange; arista light brown and minutely pubescent; fronto-orbital plate, parafacial and gena brassy pollinose; gena with few fine hairs along lower margin; facial ridge darkened; frontogenal suture black. Thorax (Fig. 17a, c): entirely gray pollinose; surfaces of dorsum of thorax and scutellum covered with conspicuous short black hairs; sternopleura, hypopleura, pteropleura, and ventral surface of abdomen yellow-white pilose; 2 katepisternal bristles; 3 postsutural supra-alar bristles, 2nd postsutural supra-alar 4X as long as 1st postsutural supra-alar; postpronotum bearing fine black hairs and anepisternum with fine yellow-white hairs; scutellum bearing two pairs of discal bristles; underside of scutellum bearing a tuft of black hairs near basal marginal bristle. Legs: coxae and femora of reddish ground color, tibiae with yellow ground color; femora covered in long black hairs interspersed among darker hairs and bristles; tarsi all black. Wings: smoky gray translucent; veins not infuscate. Abdomen (Fig. 17a): 1 pair of median marginal bristles on ST1+2, a row of marginal bristles on T3, T4 and T5; median discal bristles on T3, T4 and T5; ground color of abdomen black, with transverse bands of silver pollen on anterior half of T3, T4, and T5; underside of abdomen entirely covered in silver pollinosity. Terminalia: damage to the holotype by scavengers made the terminalia unavailable; however, other diagnostic characters are sufficient to define the species.
Female: Unknown. discal bristles on T3, anteriormost pair 1/2 the diameter of posteriormost pair; 1 pair of median discal bristles on T4 and T5; ground color of abdomen orange-brown up to tergite 5, where it darkens to brown-black; silver pollinosity covering anteriormost half of T3, T4 and T5; ST1+2 with lateral gray pollinose spots on posterior half. Terminalia (Fig. 18d, e, f): sternite 5 with two small lobes, inner margin covered in dense pollinosity appearing darker than surrounding cuticle, apical edges of lobe of sternite 5 bearing short, stout bristles interspersed with longer bristles close to apical margin; sternite 5 with a wide V-shaped median cleft, 0.46X length of sternite from lobe apex to base; cercus sharply pointed and distinctly tapered; apical section 1.7X as long as upper lobe; surstylus equilaterally oblong and scythe-like in lateral view; posterodorsal half of surstylus haired, apex with few short bristles but otherwise almost bare; tips of surstyli with a slight lobe when viewed dorsally; strongly angled inwards; 1.4X as long as cercus.
Female: no female specimen was available to us for study.

Diagnosis
Uramya sibinivora can be distinguished from all other Neotropical species of Uramya by the following combination of traits: conspicuous white pilosity covering the katepisternum, meron, and anepimeron, ventral surface of abdomen, dorsum of the thorax, scutellum white pilose, only one pair of median discal bristles on T3, and the shape of the surstylus.

Revised key to the males of Uramya Robineau-Desvoidy of Central and South America
This key builds on the work accomplished by . As observed by , females show less morphological differences between species. As a result, our key is based on male morphological characters only. For female character states, see individual species descriptions. nov.
-Three postsutural supra-alar bristles; median marginal bristles present on all tergites; one or two pairs of discal bristles on T3 and T4 6 6 Pedicel orange 7 -Pedicel dark brown 9 7 One pair of median marginals on ST1+2, T3 and T4; femora of dark ground color, covered in gray pollinosity; tibiae of yellow or dark brown ground color U. penicillata sp. nov.
-One pair of median marginals on ST1+2, an incomplete row of marginal bristles on T3, T4 and T5; legs of yellow ground color, covered in black hairs; coxae orange 8 8 Sides of thorax and underside of abdomen densely pilose; legs conspicuously yellow, thinly dark haired; T3 and T4 densely covered in erect hairs more than half as long as discal bristles; T5 lacking row of discal bristles.

U. setiventris
-Sides of thorax and underside of abdomen moderately pilose; legs densely covered in dark hair, making them appear brown; T3 and T4 densely covered in appressed hairs not more than 1/3 as long as discal bristles; row of discal bristles present on T5. Abdominal ST1+2 bearing silver pollinose spots, lacking silver pollinosity along posterior margin; wing veins strongly infuscate, giving a dark grey tonality to the leading edge of the wing.
-Abdominal ST1+2 lacking silver pollinose spots, but with a thin transverse band of silver pollinosity along posterior margin; wing veins not strongly infuscate, infuscation only visible on basal halves of R and M.
U. lunula sp. nov. Fig. 19 is a Neighbor Joining tree (NJ) (Saitou and Nei 1987) for the Uramya holotypes reared and DNA-barcoded by this inventory to date. The DNA barcode sequences obtained from the ten species of ACG Uramya displayed the strong AT bias characteristic of insect mitochondrial DNA (mean percent GC content 30.33%, SE 0.1) and displayed no insertions or deletions. Within-species variation was low compared to between-species variation. All values of DNA barcode variation were calculated within BOLD and can be re-calculated in the future as more specimens or species are added to the DNA library. 4+5 Figure 19.

Analysis
Neighbor Joining tree (NJ - Saitou and Nei 1987) comparing the species of Uramya present in ACG. Tree based on the Kimura-2-parameter (Kimura 1980) made using MEGA6 (Tamura et al. 2013) for a single specimen from each of the Uramya species discussed here. Tip labels include: species name|sample accession|species parasitized|family of parasitized caterpillar, and the image of a male in lateral view. Each sample presented here is the holotype for the species described here, except U. sibinivora. This phenogram shows the low intra-specific and high inter-specific variation of CO1 barcode sequences.