Phyllostoma Wagner, 1843: 366; part; not Phyllostoma Cuvier, 1800.

Arctibeus: Tomes, 1860: 260; incorrect subsequent spelling, not Artibeus Leach, 1821.

Vampyressa Thomas, 1900: 270; type species Phyllostoma pusillum Wagner, 1843, by original designation; described as a subgenus of Vampyrops Peters, 1865.

Vampiressa Festa, 1906: 7; incorrect subsequent spelling of Vampyressa Thomas, 1900.

Vamqyressa Cabrera, 1958: 83; incorrect subsequent spelling of Vampyressa Thomas, 1900.

Vampiressa Silva, 1975: 52; incorrect subsequent spelling of Vampyressa Thomas, 1900.

Vampuressa Coimbra, Borges, Guerra, and Mello, 1982: 34; incorrect subsequent spelling of Vampyressa Thomas, 1900.

Vampiressa Montenegro and Romero-Ruiz, 2000: 646; incorrect subsequent spelling of Vampyressa Thomas, 1900.

Vapyressa Contreras-Vega and Cadena, 2000: 287; incorrect subsequent spelling of Vampyressa Thomas, 1900.

Restricted Diagnosis of Vampyressa : The following combination of characters defines the genus Vampyressa as currently constituted (five species recognized herein): Two pairs of white facial stripes; a dorsal stripe is lacking; interfemoral membrane usually conspicuously furred (hairs may be restricted to medial area) and often fringed; a tail is lacking; inner upper incisors relatively long, bifid or weakly trifid, and usually converging distally; accessory subapical “cusp” on upper canines; caniniform p2; p4 lacks a prominent anterior cingulum; second lower molar (m2) has a well-developed, high, and bladelike entoconid; septum (medial flange of palatine processes of premaxillae) between incisive foramina perforated by large accessory medial foramen; irregularly fenestrated palate, foramen ovale relatively small and located well anterior to the caudal extension of the pterygoid (the ridge forming the anterolateral border of the eustachial groove); portions of squamosal and exoccipital bones comprising posttympanic process unfused; dental formula 2/2, 1/1, 2/2, 2/2–3 × 2 = 28–30.

Comparisons with Vampyriscus: Vampyriscus, which we recognize as a separate genus, includes three species (Vampyriscus bidens, V. brocki, and V. nymphaea). Vampyriscus can be distinguished from Vampyressa by the presence of a middorsal stripe (weakly developed in some V. nymphaea and absent in some V. brocki); lack of extensive fur and absence of a fringe of hair on interfemoral membrane; upper inner incisors long, pointed, and parallel in V. brocki, or unevenly bifid and convergent at the tips in V. bidens and V. nymphaea; normal crowns of upper canines (lack subapical cuspule); p2 not caniniform; p2 and p4 with well-developed anterior cingulae; entoconid of m2 not prominent as an elevated bladelike cusp; septum between incisive foramina narrow with a small medial accessory foramen in V. bidens, or very narrow and lacking the medial accessory foramen in V. brocki and V. nymphaea; palate comparatively flat with scattered small, porelike perforations (not obviously fenestrate as in Vampyressa); foramen ovale relatively large, adjacent to and partially occluded by high ridge of caudal extension of the pterygoid (in posterior view); bones comprising posttympanic process fused.

Vampyressa melissa Thomas, 1926: 157, type locality “Puca Tambo, Peru, altitude 7100′,” San Martín, Peru.

Vampyressa (Vampyressa) melissa: Peterson, 1968: 14; name combination.

Type Material: The holotype BMNH 26.5.3.4 consists of a relatively well-preserved skin and skull of an adult female from Puca Tambo (6°9′S, 77°16′W), Moyobamba, department of

San Martín, Peru, collected at 1480 m by R.H. Hendee (no. 408) on January 15, 1926, during the Godman-Thomas Expedition to Peru.

Measurements of the Holotype: The following measurements (mm) of the holotype are from the skin label: TL 54, HF 10.5 s.u., E 16. Additional measurements (mm) taken by one of us (V.C.T.) from the holotype are: FA 36.8, TiL 14.9, GLS 21.2, CIL 20.2, CCL 19.5, PB 5, IB 6.2, BB 8.9, BH 8.7, MB 10.4, ZB 12.5, MTRL 6.9, and MANDL 7.4.

Distribution: As defined here, Vampyressa melissa is currently known from upper-elevation localities (1180–2763 m) in Andean Colombia, Ecuador, and Peru (fig. 3).

Emended Diagnosis: A large Vampyressa (FA 34.5–37.7, n = 9; GLS 21.2–22.6, n = 9; table 3) with a broad skull (ZB 12.5–13.5, n = 8; table 3); anteriorly tapering rostrum; discrete orbitosphenoid foramina, well-developed postglenoid processes that project ventrally to the level of pterygoid processes; mesopterygoid fossa about as long as wide; large basioccipital-basisphenoid pits, and lacks a well-developed posterolabial cuspulid on p4. V. melissa is larger than V. pusilla and V. thyone, but averages smaller in most dimensions than either of the new species described below.

Redescription: Vampyressa melissa has sparse, long guard hairs conspicuous in the dorsal pelage; two pairs of pale facial stripes with the medial stripes extending above eyes from between ears to upper border of nasal horseshoe, and the lateral stripes extending below eyes from base of ears to upper corners of mouth. The pelage of the dorsum is uniformly brownish, a dorsal stripe is lacking. The noseleaf (lancet) is triangular, elongate, and with deep grooves separating the lateral flaps from the midrib; bicolored with most of lateral margins of the lancet and outer border of the horseshoe conspicuously pale yellow or cream color. The ear pinna is hairy from its base to halfway to the tip. The proximal two thirds of the forearm is densely furred.

All flight membranes are dark brown; the antebrachial membrane (propatagium) is narrow and the posterior plagiopatagium inserts on the hind limb at the base of the toe. The uropatagium is relatively narrow, its greatest breadth approximately half the length of the tibia; its posterior border has an inverted U-shape. Dorsal and ventral surfaces of the uropatagium are conspicuously haired and the posterior margin bears a paler fringe of hair that is longest toward the midline; lower legs, adjacent uropatagium, and dorsal surface of feet densely covered with reddish-brown hair.

The skull is robust; the rostrum is elevated anteriorly from a horizontal plane connecting ventral surfaces of occipital condyles and crown apices of last upper molars; premaxillae inclined posteriorly at approximately 60° from the horizontal plane (fig. 6B). The angle of the forehead with the rostrum is not abrupt (∼150°) and the forehead slopes smoothly to the rostrum in lateral outline. The ventral margin of the external nares is V-shaped and bears a small medial pit (fig. 5A, D). The external nares are emarginate dorsally (nasals comparatively short; fig. 5D). The rostrum tapers anteriorly and the postorbital constriction is narrower than interorbital breadth (fig. 4C). The sagittal crest is low, but well developed; the posterior border of the skull (inteparietal region) is not conspicuously inflated (fig. 6B). The dorsoventrally thin septum separating the triangular-shaped incisive foramina is perforated by a large accessory medial foramen (fig. 5D). The lightly and irregularly fenestrate hard palate is conspicuously domed behind the incisive foramina. The shape of the emargination of the posterior border of palate is variable, yet the mesopterygoid fossa is approximately as long as wide (figs. 4D, 8A). A discrete orbitosphenoid foramen is present. The hamular process of the pterygoid and the foramen ovale are approximately even with (on same lateral plane as) the robust postglenoid process. The paired basioccipital-basisphenoid pits are long and extend anterior to anterior margin of bullae. The caudal extension (ridge) of each pterygoid is directed posterolaterally and well separated from medium-sized bony excrescences (fig. 8A). The foramen magnum is rounded in outline.

Crowns of upper inner incisors (I1s) are large, approximately evenly bilobate, converge apically, and have well-separated roots; outer upper incisors (I2s) also are bilobate, but much smaller, extending to or slightly beyond cingulae of upper canines (fig. 5A), and anterior portions are visible in dorsal view of the skull. There are two subequal and weakly bifid lower incisors on each side of the mandible.

The upper canine (C) is about twice the length of P4, and has a conspicuous small “cusp” at the apex of the mediolateral ridge forming the lingual border of the longitudinal sulcus on its posterior crown. P3 is about a half or less the size of P4, which is nearly triangular, has a well-developed labial cingulum and a small posterior cuspule paired with a smaller cuspule on the posterolabial cingulum. The lower canine (c) has a well-developed posterior cingulum; p2 strongly resembles the lower canine, although less than half its size; p4 has a ridge along the posterior crown of the major cusp that extends across the heel of the tooth to the posterior cingulum, but lacks a discrete posterolabial cuspulid (figs. 6B, 7A).

The first upper molar (M1) has a relatively well-developed protocone, paracone, and smaller metacone. The second upper molar (M2) has a rudimentary protocone, a muchreduced metacone, and a small protoconule behind the large paracone. The first lower molar (m1) is quadrate in occlusal outline, with a large protoconid and a small blunt hypoconid. The second lower molar (m2) has a long metaconid that is twice the size of the protoconid and a large, L-shaped (bladelike) entoconid approximately the same size as the metaconid. Thomas (1926: 1) commented on the lower molars of V. melissa: “Its ... peculiarly shaped m1, and the unusual development of the characteristic cusps on m2 all readily distinguish it from the other species of the genus [sensu lato].” A small peglike m3 is present.

Comparisons: Previously, V. melissa was compared with presumed congeners Vampyriscus bidens and V. nymphaea. Goodwin (1963: 23), for example, briefly redescribed the holotype of V. melissa and commented that the skull was “about like that of Vampyriscus nymphaea but more robust.” However, these two taxa along with Vampyriscus brocki are not species of Vampyressa and need not be compared herein (see diagnosis for Vampyressa).

Vampyressa melissa is considerably larger than either of the other two known species of Vampyressa (contrast measurements of V. melissa with those of V. pusilla and V. thyone in table 3). V. melissa does merit comparison with each of the two new taxa described below (see those sections for comparisons).

During research on her doctoral dissertation, the lead author realized that Colombian specimens identified as V. melissa from upper elevations in the Andes were larger and more robust than V. melissa, and appeared to be undescribed. Here we describe this species as:

Vampyressa melissa: Lemke et al., 1982: 231.

Vampyressa melissa: Alberico et al, 2000: 55 (part).

Holotype: An adult female (FMNH 114028; figs. 4E–F, 5B, 5E, 6C, 8B, 9A) collected at Llorente (0°46′40″ N, 77°21′50″ W; 1700 m), municipio de Córdoba, departamento de Nariño, Colombia, by Kjell von Sneidern (original number: 1736) on May 10 1971. The holotype is preserved as a skin and skull (table 4).

Paratypes: An adult female (MHNUC 1514, fig. 10) caught at San Juan de Villalobos (1°33′18″ N, 76°18′19″ W; 1620 m), Vereda La Esmeralda, Santa Rosa, departamento de Cauca, Colombia, in the Caquetá River basin, eastern Colombian slopes of the eastern Andes by H.E. Ramirez-Chaves on 5 May 2005; and an adult male (IAvH 2282) collected in Parque Nacional Natural Cueva de los Guácharos (1°38′ N, 75°58′ W, 1900 m), Acevedo, departamento de Huila, Colombia, by T.O. Lemke (Lemke et al. 1982) see table 4.

Distribution: Know only from the type series from the eastern slope of the Central Cordillera (Llorente) and eastern slopes of the Eastern Cordillera (San Juan de Villalobos and Parque Nacional Cueva de los Guácharos) of the Andes in Colombia (fig. 3).

Etymology: A Quechuan word, sinchi conveys the meaning “robust and strong.” The name honors the Quechuan people of Colombia (the Ingas), and indicates the robustness of this species, the largest Vampyressa known. The name is to be treated as a noun in apposition.

Diagnosis: Vampyressa sinchi is similar to V. melissa, but larger in most dimensions with few overlapping measurements (tables 3–4, figs. 1, 2, 4E, 4F, 6C). The skull, including dentition, is larger and more robust than any other known species of Vampyressa. The strongly developed postglenoid processes extend below the level of pterygoid processes; the mesopterygoid fossa is longer than wide; the ventral surface of the postdental extension of the palate is flat, its margins comparatively sharp, and bears small winglike projections on each side at approximately the level of the orbitosphenoid foramen.

Description: Vampyressa sinchi has pale brown to dark-beige dorsal pelage; the darker dorsum contrasts with the paler venter; short, sparse guard hairs on the head and dorsum; forearm densely haired proximally for about two thirds of its length; the uropatagium is narrow (about half the length of the tibia in dried skins), furred on both surfaces, and has a sparse fringe of hairs along its entire posterior margin but longest toward the midline; tibia, adjacent uropatagium, and foot conspicuously covered with reddish-brown hairs similar to the condition in V. melissa. The third and fifth metacarpals are similar in length and the fourth metacarpal averages shorter than the third and fifth. Otherwise, V. sinchi is very similar externally to V. melissa as described in that account.

The skull of Vampyressa sinchi (figs. 4E, 4F, 6C) also is similar to V. melissa , but more robust, and larger in most dimensions with few overlapping measurements (tables 3–4). The medial, incisor-bearing anterior premaxillae of V. sinchi are inclined at approximately 60° from the horizontal plane. The external nares are emarginate dorsally; the ventral margin is V-shaped between the roots of the upper inner incisors and bears a medial pit (figs. 5B, 5E). The nasals are relatively short resulting in a comparatively longer narial opening (deeper dorsal emargination, fig. 5E). The mesethmoid plate is not swollen and its anterior edge is thin (fig. 5B).

The rostrum is elevated above the horizontal plane connecting occipital condyles and the tips of the last upper molars. The septum between the incisive foramina is dorsoventrally thin and perforated by a large accessory medial foramen (fig. 5E). The lightly and irregularly fenestrated palate is conspicuously domed behind the incisive foramina. The sagittal and lambdoidal crests are low, but well developed; the interparietal-supraoccipital region is not conspicuously inflated. The mesopterygoid fossa is longer than wide; basioccipital-basisphenoid pits are sharply defined, deep, and extend well anterior to the anterior level of the auditory bullae.

The upper inner incisors (I1s) are subequally bilobed and convergent at the tips. The dentition is similar to that of V. melissa, but more massive throughout with stronger cingular development.

Comparisons: Vampyressa sinchi is larger than all other Vampyressa known (figs. 1–2, tables 3–4; Tavares et al., 2008) and tends to have longer fur and a darker dorsum. Among species of Vampyressa, V. sinchi is most similar to V. melissa from which it differs primarily in larger size; heavier, more robust dentition; unevenly bilobed upper inner incisors, deeper dorsal emargination of anterior nares, longer than wide mesopterygoid fossa, and a distinctly larger and longer postglenoid process that extends below the level of the hamular process of the pterygoid.

Like V. melissa, V. sinchi has sloping premaxillae (approximately 60° from horizontal plane) that bear a pit located in the midline between roots of upper inner incisors; narrow anterior margin of mesethmoid plate (fig. 5E); anteriorly tapering rostrum; dorsoventrally thin septum separating wider than long incisive foramina and perforated by a large accessory medial foramen; conspicuously domed hard palate behind incisive foramina; orbitosphenoid foramina (fig. 9A); foramen ovale and ventral projection of hamular process of pterygoid on same lateral plane as the base of the postglenoid process (fig. 8B); large basisphenoid-basioccipital pits that extend well anterior to anterior surfaces of auditory bullae; caudal ridge from pterygoid well separated from comparatively small boney excrescences (located on basisphenoid anteromedial to auditory bullae) (fig. 8B); rounded foramen magnum; robust mandibular rami and angular process. Like V. melissa, in V. sinchi the outer upper incisors are visible from above; p4 has a posterior ridge that extends down from the crown of the major cuspid and across heel to the posterior cingulum, and p4 also lacks a posterolabial cuspulid. V. sinchi and V. melissa are the only species of Vampyressa that retain the spiculelike last lower molar; the dental formula is 2/2, 1/1, 2/2, 2/3 × 2 = 30.

The late Charles O. Handley, Jr., acquired four specimens of a large Vampyressa from the Bocas del Toro region in the 1960s during fieldwork in Panama related to a countrywide survey of the ectoparasites of vertebrates, primarily mammals. Although cataloged as Vampyressa melissa, Handley (1966: 767) reported these specimens as “Vampyressa species,” but never got around to describing them. An additional specimen from the same region was acquired by Robert K. Ender in 1976. These specimens are described here as:

Holotype: An adult male (USNM 319283; figs. 4A-B, 6A) caught in evergreen forests of Rancho Mojica, upper Changena River, ±20 miles [32.2 km] SSW of Changuinola (9°6′ N, 82°34′ W; 1463 m), Provincia Bocas del Toro, Panama, by Vernon J. Tipton (original number 8006) on 13 September 1961. The holotype is preserved as a skin and skull (table 5).

Paratypes: The skins and skulls of one adult female (USNM 319282; figs. 8C, 9B) and two adult males (USNM 319284 [figs. 5F, 7B], 319285 [fig. 5C]) caught at Rancho Mojica, upper Changena River, ±20 miles SSW of Changuinola (09°6′ N, 082°34′ W, 1463 m), Provincia Bocas del Toro, Panama (original numbers V.J. Tipton 8006, 8120, 8122). A fluid-preserved specimen of a male (USNM 520851), with skull removed, collected by Robert K. Enders at “Fish Camp” (08°58′N, 082°40′W, 1494 m), Bocas del Toro, Panama on 26 March 1976.

Distribution: Know only from provincia de Bocas del Toro, Panama (fig. 3).

Etymology: The name elisabethae honors the legacy of the late Elisabeth Klara Victoria Kalko in the study of the natural history, ecology, conservation, vocal recognition, and behavior of bats.

Diagnosis: A large Vampyressa (FA 36.2–37.8, n = 5; GLS 22.9–23.0, n = 5; table 5) having a relatively long, narrow skull (ZB 12.0–12.6, n = 5; table 5) with nearly parallel sides of the rostrum. Externally and dentally similar in appearance and size to V. melissa and V. sinchi, V. elisa- bethae has a swollen anterior margin of mesethmoid plate (fig. 5C); dorsal contour of external nares not deeply emarginate (fig. 5F); an orbitosphenoid fissure (instead of a discrete foramen, figs. 4B, 9B); caudal border of pterygoid extending posteriorly as a low ridge over basisphenoid to near contact with the large bony excrescence (figs. 8C, 9B); laterally ovoid foramen magnum; unequally bilobate to weakly trilobate upper inner incisors (fig. 5C); a small cuspulid on posterolabial crown of p4 (fig. 7B); m3 absent; dental formula 2/2, 1/1, 2/2, 2/2 × 2 = 28.

Description: Vampyressa elisabethae is closely similar in color pattern and other external features to both V. melissa and V. sinchi, although slightly smaller than V. sinchi and averaging equal to or slightly larger than V. melissa. On average, V. elisabethae has slightly paler dorsal pelage (pale ochre and yellowish brown to beige) than do either V. melissa or V. sinchi. The forearm is densely furred for a little more that its proximal half, and dorsal and ventral surfaces of the propatagium and plagiopatagium are hairy along sides of the body. The dorsal and ventral uropatagium, legs, and feet are sparsely furred, except toward the ventral midline of the uropatagium, where the fur is especially dense and long. The uropatagium lacks a conspicuous fringe of hair along the entire posterior margin. Like V. melissa and V. sinchi, the uropatagium is relatively narrow, its greatest breadth measuring less than half the length of the tibia. The third and fourth metacarpals are subequal, and the fifth metacarpal is longer than either the third or fourth. The tibia is short (12.3–12.9, n = 5) relative to the overall size of the bat.

Vampyressa elisabethae is cranially and dentally similar to V. melissa and V. sinchi, but can be distinguished by its relatively longer, narrower skull; nearly parallel sides of the rostrum; nearly straight profile of frontals and rostrum in lateral view (fig. 6A); less convergent upper toothrow; and narrower zygomatic breadth (ZB 12.0–12.6, n = 5). The anterior plane of premaxillae is near vertical (80°) from the horizontal plane of skull; the rostrum is aligned with the cranium (tips of canines and second upper molars on same horizontal plane as ventral surfaces of occipital condyles); lower margin of anterior external nares U-shaped and lacking a medial pit between roots of upper inner incisors; anterior margin of mesethmoid plate swollen (fig. 5C); dorsal contour of external nares not deeply emarginate (fig. 5F); an orbitosphenoid fissure present (figs. 4B, 9B); incisive foramina comparatively round with medial septum dorsoventrally thick and perforated by a comparatively small accessory medial foramen; palate not conspicuously domed behind incisive foramina; short, shallow basioccipital-basisphenoid pits; caudal border of pterygoid extending posteriorly as a low ridge to near contact with large bony excrescence on basisphenoid; foramen magnum laterally ovoid (fig. 8C); ventral surface of postdental palate weakly depressed (indented) along ventral midline and its lateral margins rounded along at least part of their length; foramen ovale and pterygoid process located behind medial base of postglenoid process; each dentary is proportionally slender, dorsoventrally narrow, has a low coronoid process, and a small articular process that is close to the small angular process; upper toothrows not strongly convergent anteriorly; unequally bilobate to weakly trilobate upper inner incisors; upper outer incisors not easily visible in dorsal view of the skull; a posterolabial cuspulid present on p4, but lacking a medial ridge from major anterior cuspid to cingulum on heel; m3 absent; the dental formula is 2/2, 1/1, 2/2, 2/2 × 2 = 28.

Comparisons: Vampyressa elisabethae needs no comparison with the much smaller V. pusilla and V. thyone. Although externally similar to V. melissa and V. sinchi, V. elisabethae has somewhat paler pelage, less hairy feet and uropatagium (except ventrally near midline), and shorter tibiae. The skull is conspicuously narrower and the anterior premaxillae more vertically oriented (near 80° from the horizontal plane versus approximately 60° in V. melissa and V. sinchi). The anterior nasal cavity is considerably different in that the ventral margin in V. elisabethae is U-shaped and lacks a medial pit (V-shaped with a medial pit between roots of upper inner incisors in the other two taxa), less emarginate upper border (deeper emargination in V. melissa and the much more deeply emarginate border in V. sinchi; fig. 5), and swollen anterior margin of the conspicuously thicker mesethmoid plate (plate thinner in V. melissa and V. sinchi; fig. 5). The sphenorbital opening in the cranium is a fissure in V. elisabethae in contrast with all other species of Vampyressa in which the opening is a foramen. The morphology of the anterior palate differs between V. elisabethae, V. melissa, and V. sinchi (fig. 5) and the palate is domed behind the incisive foramina in V. melissa and V. sinchi, but not domed in V. elisabethae. The foramen magnum is transversely ovoid in V. elisabethae, but more rounded in all other Vampyressa. Each dentary is massive, the coronoid process high, and both articular and angular processes are larger and not particularly close together in V. melissa and V. sinchi (fig. 6B–C), whereas in V. elisabethae each dentary is proportionally more slender, dorsoventrally narrow with a low coronoid process, and the smaller articular process is closer to the smaller angular process (fig. 6A).

Vampyressa melissa and V. sinchi obviously are closely related as reflected by their close similarity. V. elisabethae is not as closely related to these two species, nor is it closely related to V. pusilla and V. thyone based on its clearly distinctive features. Nevertheless, all known species of Vampyressa share the same major dental features, particularly the caniniform p2 and the characteristic well-developed, high, and bladelike entoconid on m2.