Peropteryx pallidoptera, Lim & Engstrom & Reid & Simmons & Voss & Fleck, 2010

Lim, Burton K., Engstrom, Mark D., Reid, Fiona A., Simmons, Nancy B., Voss, Robert S. & Fleck, David W., 2010, A New Species of Peropteryx (Chiroptera: Emballonuridae) from Western Amazonia with Comments on Phylogenetic Relationships within the Genus, American Museum Novitates 3686, pp. 1-20 : 3-14

Peropteryx pallidoptera , new species

Pale-winged Doglike Bat

Figures 1 , 2 , 3

HOLOTYPE: An adult male ( ROM 104396 ) collected 66 km S of Pompeya Sur (00 ° 48 9 S, 76 ° 24 9 W), Orellana Province, Ecuador by F.A. Reid, and F. Sornoza (field number F37227) on 8 May 1995. The holotype was prepared as a skin, skull, and partial skeleton.

PARATYPES: An adult male ( AMNH 74107 ) prepared as a skin and skull collected at Orosa (03 ° 26 9 S, 72 ° 08 9 W), Amazon River , Loreto, Peru, by Alfonso and Ramón Olalla on 28 November 1926. Seventeen individuals prepared as fluid-preserved specimens collected at Nuevo San Juan (05 ° 14 9 50 0 S, 73 ° 09 9 50 0 W, 150 m above sea level), Río Gálvez, Loreto, Peru, by Robert S. Voss and David W. Fleck in 1998 and 1999, including 1 adult male ( MUSM 13226 ) , 14 adult females ( AMNH 272671 , AMNH 272726 , AMNH 272854 , AMNH 272855 , AMNH 272827 , AMNH 273042 , AMNH 273116 , AMNH 273185 , MUSM 13228 , MUSM 13230 , MUSM 15246 , MUSM 15249 , MUSM 15250 , MUSM 15252 ) and 2 subadult females ( MUSM 13227 , MUSM 13229 ) .

DISTRIBUTION: The new species is currently known from just three localities in primary lowland forests of eastern Ecuador and northern Peru at elevations, 400 m above sea level. It is found sympatrically with P. kappleri and P. leucoptera at Nuevo San Juan. The new species and P. macrotis have overlapping distributions in the western Amazon but their closest points of documented occurrence are approximately 50 km apart (between Orosa and Puerto Indiana on the Amazon River of Peru).

ETYMOLOGY: The specific epithet refers to the pale wings and is compounded from the Latin words pallidus (‘‘pale’’) and ptera (‘‘wing’’).

MEASUREMENTS: External measurements are presented in table 1 and cranial measurements are presented in table 2 for Peropteryx pallidoptera and other congeneric species.

DIAGNOSIS: Superficially, Peropteryx pallidoptera appears most similar to P. leucoptera because they both have transparent wings. However, the new species has paler brown arms and digits (dark brown in P. leucoptera ), the skin from the wing tip to the elbow is translucent and evenly colored with a tinge of brown (gradually darkens in color from wing tip to body in P. leucoptera ; fig. 1), the ears are not connected by a band of skin, the rostrum of the skull is not broad, and the lateral pterygoid pits are not as large and deep as they are in P. leucoptera (fig. 2). In contrast, P. pallidoptera is morphologically similar to P. kappleri , P. macrotis , and P. trinitatis with the ears separated, rostrum of the skull not broad, and the lateral pterygoid pits small and shallow (fig. 2). However, the new species does not have dark brown wings and digits, the rostrum is not inflated dorsally, and the posterior border of the mesopterygoid extension is not level with the anterior border of the basisphenoid and lateral pterygoid pits, all of which are characters seen in P. kappleri , P. macrotis , and P. trinitatis . Furthermore, P. pallidoptera has a tiny peglike upper anterior premolar similar to that of P. trinitatis but different from those of P. kappleri and P. macrotis ; the latter two species have a slightly larger tooth, bearing a distinct posterior cusp. Peropteryx kappleri is the largest species in the genus with no overlap in forearm length and cranial measurements (except postorbital constriction) with the other species ( tables 1 and 2). In general, P. pallidoptera is smaller in cranial size than P. leucoptera , P. macrotis , and P. kappleri , but it is larger than P. trinitatis . This size gradation is most evident in same-sex comparisons of condyloincisive length ( table 3 ).

DESCRIPTION: The new species is a medium-sized Peropteryx ( table 2 ). The dorsal fur is relatively long (approximately 8 mm) and is a uniform medium brown with slightly paler hair bases. The ventral fur is similar but slightly lighter in color. The ears are brown and are not in contact with each other, although they are close together on the forehead. The interfemoral membrane is brown, as is the wing membrane from the body to the elbow, but the distal portion is translucent with a tinge of brown (fig. 1). The arms and digits of the wings are medium brown. There is a poorly developed wing sac situated on the leading edge of the propatagium with an outward lateral opening.

The skull of P. pallidoptera has slender postorbital processes and a relatively narrow rostrum, but the rostrum is not dorsally inflated. The relatively shallow, undivided basisphenoid pit has two small anterolateral pterygoid pits and the mesopterygoid extension protrudes posteriorly into the basisphenoid region (fig. 2). The upper anterior premolar is tiny and peglike. The dental formula is i 1/3, c 1/1, p 2/2, m 3/3 with a total of 32 teeth.

COMMENTS: The cranial measurements for males of P. pallidoptera are all within the observed range of variation for females ( table 2 ), which suggests that there is little or no sexual dimorphism in skull size. However, although the sample size for male P. pallidoptera is small (N 5 2), males are smaller than females with no overlap in forearm length and weight ( table 1 ). In contrast, females of P. macrotis are significantly larger than males (p, 0.02) for all cranial measurements except breadth of braincase and postorbital constriction. Similarly, females of P. macrotis are significantly larger than males (p, 0.05) for all external measurements except length of tail and length of tragus. Brosset and Charles- Dominique (1990) did not observe sexual dimorphism for P. trinitatis but they only compared 1 male and 1 female. Although our sample size was not much better (at 2 males and 2 females), females were larger for all cranial measurements except postorbital constriction and there was no overlap in range ( table 2 ). The male P. trinitatis reported by Brosset and Charles-Dominique (1990) was large and similar to our females except for maxillary toothrow length. However, with larger sample sizes for external measurements (10 males and 10 females), females of P. trinitatis are significantly larger (p, 0.003) than males except for length of tail and length of tragus. The sample sizes for P. kappleri and P. leucoptera were too small for making meaningful comment on cranial sexual size dimorphism. External measurements suggest that females of P. kappleri are larger than males (p, 0.005 for total length, forearm length, and weight), but there is no indication of sexual dimorphism in any of the external measurements for P. leucoptera .

NATURAL HISTORY

ECUADOR: Peropteryx pallidoptera was one of 66 species of bats that we captured in Block 16 in 1995 and 1996 ( Reid et al., 2000). The holotype was caught in a mist net set at the entrance to a small cave next to a muddy, spring-fed saladero (mineral-rich seep) that periodically flooded the surrounding forest. The cave appeared to have been used primarily as a night roost, because few bats were found to occupy it in the daytime.

Another six specimens of Peropteryx pallidoptera (easily recognized as such by their transparent wing tips) were captured and released by Engstrom, who netted them over a larger saladero (at least 1 km in diameter) located near the edge of the Tiputini River about 10 minutes by boat north of the Tiputini Biodiversity Station (00 ° 38 9 S, 76 ° 09 9 W). The small stream that fed this saladero tumbled over the edge of a cliff ca. 10 m high, and a shallow cave was present behind the waterfall. The bats apparently used the cave as a diurnal roost: Engstrom saw small, brown emballonurids in the cave before it got dark, and he caught the six P. pallidoptera in the net nearest the cave exit at dusk.

PERU: We recorded a total of 58 species of bats at Nuevo San Juan in 1998 and 1999 ( Fleck et al., 2002). Of 503 recorded bat captures at this locality, 372 were in mist nets and 311 were made at diurnal roosts. All 17 specimens of Peropteryx pallidoptera from Nuevo San Juan were taken from roosts. In total, we found 10 roosts occupied by this species in a variety of circumstances as described below (numbers refer to entries in a field catalog of bat roosts).

Roost 2: Dark cavities among the roots and buttresses of a fallen tree in well-drained primary forest. Exactly where the bats were roosting is unknown because they had been spooked by kids who were shooting at them with arrows. A total of eight bats were collected as they perched on nearby tree trunks. Three (one adult male, one adult female, and one subadult female) were Peropteryx pallidoptera and five were Saccopteryx bilineata ; an unknown number of either or both species might have escaped (22 May 1998).

Roost 6: A half-open hollow log (about 50 cm inside diameter) in old secondary growth near a stream. Two bats were found roosting here, only one of which (an adult female Peropteryx pallidoptera ) was collected (11 June 1998).

Roost 15: An undercut earth bank in the side of a stream gully in well-drained primary forest. Two adult female and one subadult female Peropteryx pallidoptera occupied this roost, where they were found hanging well apart from one another (30 June 1998).

Roost 20: A small dark cavity of roots and earth at the base of a large fallen tree in welldrained primary forest. Two bats were found roosting here about 50 cm above the ground, of which only one (an adult female Peropteryx pallidoptera ) was collected (8 July 1998).

Roost 22: A dark horizontal chamber under a buttress of a fallen tree in well-drained primary forest. Two adult female Peropteryx pallidoptera were found roosting here, about 80 cm above the ground (8 July 1998).

Roost 31: A deep armadillo burrow in the side of a stream headwater gully in welldrained primary forest. One adult female Peropteryx pallidoptera was found roosting here in the company of at least two Micronycteris matses and three Carollia brevicauda . No bats were seen to escape this roost by flying out of it, but some might have escaped by flying deeper inside (4 September 1999).

Roost 56: The unmodified leaf of a wild banana plant in hilltop primary forest. The single bat (a pregnant female Peropteryx pallidoptera ) observed and collected here was hanging from the middle of the leaf about 1 m above the ground (16 September 1999).

Roost 73: A hole in the ground in the side of a stream headwater gully on a hillside in well-drained primary forest. Two bats were observed roosting here, of which one (a pregnant female Peropteryx pallidoptera ) was collected (22 September 1999).

Roost 77: A hole in the side of a stream headwater gully in hilltop primary forest. Four bats were observed using this roost, of which two were collected. One of those collected was an adult female Peropteryx pallidoptera , and the other was an adult male P. leucoptera (23 September 1999).

Roost 102: The undercut bank of a small stream in well-drained primary forest. Only a single bat (an adult female Peropteryx pallidoptera ) was observed and collected here (11 October 1999).

Roost 147: The ‘‘underside of a log’’ [as described by the Matses collector; presumably elevated somehow above the ground] in well-drained primary forest in a valley bottom. A single lactating adult female Peropteryx pallidoptera was collected here, but it seems likely that one or more nursing young may have been present but unobserved (27 October 1999).

Roost 154: The underside of a fallen tree in primary upland forest near a small stream. A single female Peropteryx pallidoptera was in the roost (2 November 1999).

The specimen from Orosa was collected by Alfonso and Ramón Olalla, members of an Ecuadorean family of professional collectors employed by the American Museum of Natural History, who worked at or near the mouth of the Río Orosa on the right (south) bank of the Amazon from 30 August to 11 December 1926. According to Wiley (in press), the Olallas’ camp at Orosa must have been in the floodplain of the Amazon, which is so wide at this point (about 15 km) that most of their specimens certainly came from floodplain habitats. Although the forest within several kilometers of the riverbank at the Olallas’ presumed campsite (ca. 3 ° 31 9 31 0 S, 73 ° 11 9 22 0 W; Wiley, in press) is seasonally flooded varzea, the river was probably at or near its lowest annual stage in August– December, so this habitat would have been easy to traverse on foot.

Figures

Fig. 1. The contrasting color patterns of the wings spread on the left and closed on the right for three species of Peropteryx: top, P. leucoptera (ROM 112531) with white wing tips, brown toward the elbow and brown digits; middle, P. pallidoptera (ROM 104396) with uniform white from the wing tips to the elbow and light brown digits; and bottom, P. macrotis (ROM 108523) with uniform brown from the wing tips to the elbow and brown digits.

Fig. 2. Differences in basisphenoid pit and lateral pterygoid pits at the anterior end (toward the top) for three species of Peropteryx: top, P. leucoptera (ROM 112531) with large pterygoid pits separated by a mesopterygoid extension; middle, P. pallidoptera (ROM 104396) with small pterygoid pits separated by a mesopterygoid extension; and bottom, P. macrotis (ROM 108467) with small pterygoid pits separated by the basisphenoid pit.

Fig. 3. Skull of the holotype of Peropteryx pallidoptera (ROM 104396) with dorsal, ventral, and lateral views of the cranium, and lateral view of the left mandible from top to bottom. The greatest length of the skull is 14.1 mm.

Tables

TABLE 1 External measurements (in mm) and weight (in g) for adult specimens of Peropteryx The average and range of measurements for each sex of the five species of Peropteryx and the probability (*, 0.05) of sexual dimorphism within each species with sample sizes.2 are summarized in boldface. Abbreviations of measurements are listed in the Materials and Methods section, and localities of specimens examined are listed in appendix 1.

Species	Specimen	Sex	TL	TV	HF	EAR	TR	FA	WT	Country
P. pallidoptera	ROM 104396	male	57	15	9	15	5	40	4	Ecuador
P. pallidoptera	MUSM 13226	male	62	11	9	15	—	39	4.3	Peru
			59.5	13	9	15	5	39.5	4.15
			57–62	11–15	9	15	5	39–40	4–4.3
P. pallidoptera	AMNH 272671	female	65	11	9	15	—	41	5.2	Peru
P. pallidoptera	AMNH 272726	female	65	13	9	15	—	41	5.6	Peru
P. pallidoptera	AMNH 272854	female	59	14	9	15	—	—	5.5	Peru
P. pallidoptera	AMNH 272855	female	60	13	8	15	—	—	5.0	Peru
P. pallidoptera	AMNH 272827	female	65	14	10	15	—	42	5.2	Peru
P. pallidoptera	AMNH 273042	female	62	12	9	15	—	41	5.2	Peru
P. pallidoptera	AMNH 273116	female	64	13	10	16	—	43	7.1 1	Peru
P. pallidoptera	AMNH 273185	female	66	12	10	16	—	42	6.0	Peru
P. pallidoptera	MUSM 13228	female	61	13	10	15	—	43	4.5	Peru
P. pallidoptera	MUSM 13230	female	58	13	9	15	—	—	4.9	Peru
P. pallidoptera	MUSM 15246	female	61	13	9	16	—	—	5.9	Peru
P. pallidoptera	MUSM 15249	female	63	12	9	17	—	42	6.7 1	Peru
P. pallidoptera	MUSM 15250	female	67	14	9	16	—	43	7.6 1	Peru
P. pallidoptera	MUSM 15252	female	66	11	10	15	—	43	5.2	Peru
			63.0	12.7	9.3	15.4	—	42.1	5.3
			58–67	11–14	8–10	15–17	—	41–43	4.5–6.0
			—	—	—	—	—	—	—
P. l. leucoptera	AMNH 273182	male	69.0	13.0	10.0	19.0	—	45.0	8.0	Peru
P. l. leucoptera	AMNH 273197	male	66.0	16.0	10.0	20.0	—	45.0	8.5	Peru
P. l. leucoptera	MUSM 15247	male	66.0	16.0	10.0	20.0	—	46.0	8.1	Peru
P. l. leucoptera	MUSM 15251	male	69.0	13.0	9.0	19.0	—	46.0	8.5	Peru
P. l. leucoptera	AMNH 267838	male	63.0	13.5	8.0	16.5	—	41.0	6.4	French Guiana
P. l. leucoptera	AMNH 266012	male	63.0	13.0	9.0	16.0	—	43.0	6.0	French Guiana
P. l. leucoptera	AMNH 267280	male	69.0	15.0	8.0	16.0	—	43.0	6.1	French Guiana
P. l. leucoptera	AMNH 267088	male	65.0	14.0	9.0	15.0	—	42.0	6.2	French Guiana
P. l. leucoptera	ROM 41530	male	56	5	8	16	7	43	—	Guyana
P. l. leucoptera	ROM 107458	male	61	11	9	16	5	44	6	Guyana
P. l. leucoptera	ROM 112530	male	57	10	9	16	6	42	4	Guyana
P. l. leucoptera	ROM 112531	male	60	11	9	16	6	42	5	Guyana

TABLE 2 Cranial measurements (in mm) for adult specimens of Peropteryx The average and range of measurements for each sex of the five species of Peropteryx and the probability (*, 0.05) of sexual dimorphism within P. macrotis (other species have small sample sizes) are summarized in boldface. Abbreviations of measurements are listed in the Materials and Methods section, and localities of specimens examined are listed in appendix 1.

Species	Specimen	Sex	GLS	CIL	ZB	MB	BBC	POC	CM3	M3M3
P. pallidoptera	ROM 104396	male	14.1	12.5	8.2	7.3	6.4	2.8	5.3	5.9
P. pallidoptera	AMNH 74107	male	—	—	8.1	7.2	6.5	2.9	5.0	5.9
P. pallidoptera	MUSM 13226	male	—	—	—	—	—	2.6	5.2	5.8
			14.1	12.5	8.2	7.3	6.5	2.8	5.2	5.9
			14.1	12.5	8.1–8.2	7.2–7.3	6.4–6.5	2.6–2.9	5.0–5.3	5.8–5.9
P. pallidoptera	AMNH 272671	female	14.1	12.8	8.3	7.1	6.5	2.6	5.3	6.1
P. pallidoptera	AMNH 272726	female	—	—	8.4	7.3	6.7	2.9	5.5	6.1
P. pallidoptera	AMNH 272827	female	14.1	12.7	8.7	7.4	6.6	2.9	5.3	6.4
P. pallidoptera	MUSM 13227	female	13.7	12.2	8.0	6.9	6.6	2.7	5.1	5.8
P. pallidoptera	MUSM 13228	female	13.8	12.5	8.1	7.2	6.5	2.6	5.2	6.1
P. pallidoptera	MUSM 13229	female	13.9	12.7	8.3	7.2	6.4	2.8	5.5	6.1
P. pallidoptera	MUSM 13230	female	13.6	12.4	8.1	7.1	6.3	2.6	5.0	5.7
			13.9	12.6	8.3	7.2	6.5	2.7	5.3	6.0
			13.6–14.1	12.2–12.8	8.0–8.7	6.9–7.4	6.3–6.7	2.6–2.9	5.0–5.5	5.7–6.4
P. l. leucoptera	ROM 41530	male			9.5	8.0	7.2	3.3	6.2	7.0
P. l. leucoptera	ROM 107458	male	15.6	14.1	9.5	7.9	7.3	3.1	6.1	6.9
P. l. leucoptera	ROM 112531	male	15.5	14.1	9.6	7.8	7.2	3.3	6.1	6.7
P. l. leucoptera	ROM 113612	male	14.9	13.9	9.3	7.6	7.0	3.3	6.1	6.8
			15.3	14.0	9.5	7.8	7.2	3.3	6.1	6.9
			14.9–15.6	13.9–14.1	9.3–9.6	7.6–8.0	7.0–7.3	3.1–3.3	6.1–6.2	6.7–7.0
P. l. cyclops	BM 24.3.1.6 1	male	16.2 2	14.5 3	—	9.2	7.8	3.2	6.5	—
P. kappleri	ROM 78055	male	—	—	9.9	8.4	7.7	2.9	6.6	7.5
P. kappleri	ROM 101123	male	18.2	16.4	10.3	8.6	7.7	2.8	7.2	7.8
			18.2	16.4	10.1	8.5	7.7	2.9	6.9	7.7
			18.2	16.4	9.9–10.3	8.4–8.6	7.7	2.8–2.9	6.6–7.2	7.5–7.8
P. kappleri	AMNH 272798	female	17.5	16.2	10.4	8.5	7.5	3	7.1	7.7
P. kappleri	ROM 100910	female	17.5	16.3	10.2	8.3	7.4	2.8	7.5	8.1
			17.5	16.3	10.3	8.4	7.5	2.9	7.3	7.9
			17.5	16.2–16.3	10.2–10.4	8.3–8.5	7.4–7.5	2.8–3.0	7.1–7.5	7.7–8.1
P. trinitatis	ROM 107822	male	12.4	11.2	7.2	6.7	6	2.8	4.4	5.1

TABLE 3 Morphological comparison among the five species of Peropteryx

	P. kappleri	P. leucoptera	P. macrotis	P. pallidoptera	P. trinitatis
Wings	Uniformly brown	Translucent, gradually	Uniformly brown	Translucent and evenly	Uniformly brown
		darkening to brown from		tinged pale brown
		wing tips to body
Arms and digits	Dark brown	Dark brown	Dark brown	Medium brown	Dark brown
Ears	Not connected	Connected	Not connected	Not connected	Not connected
Rostrum of skull	Not broad	Broad	Not broad	Not broad	Not broad
	Dorsally inflated	Not dorsally inflated	Dorsally inflated	Not dorsally inflated	Dorsally inflated
Lateral pterygoid pits	Small and shallow	Large and deep	Small and shallow	Small and shallow	Small and shallow
Mesopterygoid posterior	Does not extend	Extends	Does not extend	Extends	Does not extend
extension into
basisphenoid region
Upper anterior premolar	With posterior cusp	Peglike	With posterior cusp	Peglike	Peglike
Overall size	Large	Medium	Small	Small	Small
	Forearm 47–54	Forearm 41–46	Forearm 39–45	Forearm 39–43	Forearm 41.9
Condyloincisive length	16.4 (males, N 5 1)	13.9–14.1 (males, N 5 3)	12.7–13.7 (males,	12.5 (males, N 5 1)	11.2–11.6 (males,
			N 5 5)		N 5 2)
	16.2–16.3 (females,		13.6–14.6 (females,	12.2–12.8 (females,	11.8–12.2 (females,
	N 5 2)		N 5 9)	N 5 6)	N 5 2)