First Records of 10 Bat Species in Guyana and Comments on Diversity of Bats in Iwokrama Forest

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Acta Chiropterologica, l(2): 179-190,1999 PL ISSN 1508-1 109 O Museum and Institute of Zoology PAS

First records of 10 bat species in Guyana and comments on diversity of bats in Iwokrama Forest

BURTONK. LIM', MARKD. ENGSTROM~,ROBERT M. TIMM~,ROBERT P. ANDERSON~, and L. CYNTHIAWATSON~

'Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen's Park, Toronto, Ontario M5S 2C6, Canada; E-mail: [email protected] 2Natural History Museum and Department of Ecology & Evolutionary Biology, University of Kansas, Lawrence, Kansas 66045-2454, USA 3Centrefor the Study of Biological Diversity, University of Guyana, Turkeyen Campus, East Coast Demerara, Guyana

Ten species of bats (Centronycteris-maximiliani,Diclidurus albus, D. ingens, D. isabellus, Peropteryx leucoptera, Micronycteris brosseti, M. microtis, Tonatia carrikeri, Lasiurus atratus, and Myotis riparius) collected in the Iwokrarna International Rain Forest Programme site represent the first records of these taxa from Guyana. This report brings the known bat fauna of Guyana to 107 species and the fauna of Iwokrama Forest to 74 species. Measurements, reproductive data, and comments on taxonomy and distribution are provided.

Key words: Chiroptera, Neotropics, Guyana, Iwokrama Forest, inventory, species diversity

on the first of two field trips that constituted the mammal portion of the faunal survey for The mammalian fauna of Guyana is Iwokrama Forest coordinated through The poorly documented in comparison with Academy of Natural Sciences of Philadel- neighbouring countries in northern South phia. Records from previously unreported America. Most of its species and their distri- specimens at the Royal Ontario Museum are butions are inferred (e.g., Eisenberg, 1989) also presented to augment distributional data. from mammals reported in Venezuela (e.g ., Handley, 1976) and Suriname (e.g., Husson, MATERIALSAND METHODS 1978). Two recent publications (Parker et al., 1993; Smith and Kerry, 1996) reported on Study Sites separate, small single-site collections from Fieldwork was conducted primarily from 9 March Guyana and indicate a renewed interest in to 12 April 1997 in the Iwokrama International Rain the diversity of Guyanese mammals. Herein, Forest Programme site in central Guyana (Fig. I). we record 10 additional species of bats for Iwokrama Forest encompasses 3,600 krn2 of primary the country and provide comments on taxon- mixed lowland tropical rainforest set aside by the government of Guyana in 1990 under the auspices of omy, distribution, measurements, and repro- the Commonwealth Secretariat for sustainable man- ductive data. This report is based primarily agement and conservation of biodiversity (Hawkes and 180 B. K. Lim, M. D. Engstrom, R. M. Timm, R. P. Anderson, and L. C. Watson

forest with a clearing and road in the vicinity. Collec- tions were made on 29 March and 12 April 1997. Atlantic Before the access road was constructed and Ocean Iwokrama Forest was established, collections were made on 4,5, 10, and 11 October 1990,30 km NE of Surama in mixed lowland forest in what is now the south-central portion of Iwokrama Forest. From 17 to 31 July 1994 collections were made at Surama Saw- mill, 5 km SE of Surama adjacent to the southwestern extent of Iwokrama Forest in mixed lowland forest, which recently had been selectively logged. Additional records from previously unreported specimens at the ROM, which augment distributional t information for the species recorded herein, are also lwokrama presented (Fig. 1). These include collections made on 26 October and from 21 to 24 November 1996 at Gunn's Strip in the Upper Takutu-Upper Essequibo Region in southern Guyana. Gunn's Strip is a Wai- Wai village situated in seasonally flooded riparian forest bordering interior lowland shrub savanna. Other collections were made on the upper Essequibo River about 7 km S of Gunn's Strip from 15 to 20 November FIG. 1. Map of Guyana showing Iwokrama Forest and 1996. The habitat there was tall evergreen hill-land the collecting sites of the 10 new records of bat forest. Collections were made from 28 October to 6 species for the country reported in this paper: (I) November 1996 at Chodikar River, about 55 km SW Georgetown; (2) Bada Creek; (3) Pakatau Falls; (4) of Gunn's Strip near the border with Brazil in tall Clearwater Camp; (5) Iwokrama Field Station, evergreen non-flooded hill-land forest. Other records Kurupukari; (6) Three Mile Camp; (7) 30 km NE of were derived from specimens made by other collectors Surania; (8) Surama Sawmill; (9) Illia Wau River; (10) and are described in the species accounts. Gunn's Strip; (11) 7 km S of Gunn's Strip; and (12) The primary method used to capture bats was Chodikar River standard-weight (70-denier) mist nets of various sizes set in different microhabitats usually opened from Wall, 1993). One all-season access road, approxi- dusk to midnight. Bats flying below 3 m in the forest mately 70 km in length, bisects Iwokrama Forest from understorey were caught in regular-sized mist nets the northeast near Kurupukari on the Essequibo River (2.6x6.0 m or 2.6x12.0 m) typically set across flyways to the southwest near Surama, bordering the North such as streams and trails or near clearings and fruiting Rupununi Savannas. trees. Bats flying upwards to 20 m in the forest canopy The collecting sites (Fig. 1; see also Gazetteer) were caught with large-sized mist nets (9.1x30.5 m) within Iwokrama Forest included: (I) Pakatau Falls is set over roads or rivers within rainforest. Mist netting on the Siparuni River near the mouth of Pakatau Creek was supplemented with 4-bank harp traps usually set in the northwest in mixed, poorly drained forest near across narrow streams or trails, and shotguns (12 high hills dominated by Mora, Pentaclethra, and gauge and 0.410 calibre) with ammunition ranging Licania. Collections were made from 9 to 16 March from number 6 to 12 shot. Although the searching for 1997; (2) Three Mile Camp is in the northeast along roost sites was not an active part of our field protocol, the road approximately 5 km southwest of Kurupukari we checked potential roosts when encountered. in undulating terrain on well-drained brown sand with lowland forest dominated by Licania, Chlorocardium, Specimens and Measurements and Eschweilera. Collections were made from 18 to 28 March 1997; (3) Cleanvater Camp along the Burro Specimens were prepared as dried skins with Burro River approximately 25 km WNW of Kuru- skeletons or fixed in 10% formalin with storage in pukari is in the north in mixed lowland forest domi- 70% ethanol. All specimens had tissue samples (heart, nated by Chlorocardium, Eperua, and Eschweilera on kidney, spleen, liver) frozen initially in liquid nitrogen well-drained white sand. Collections were made from with later storage in an ultracold freezer (-80°C). 30 March to 11 April 1997; (4) Iwokrama Field Voucher specimens were deposited at the Angelo State Station is in the northeast on the Essequibo River Natural History Collection (ASNHC), Royal Ontario about 1 km N of Kurupukari in disturbed alluvial Museum (ROM), U.S. National Museum of Natural First records of 10 species of bats in Guyana 181

History (USNM), University of Guyana, Centre for the basisphenoid region) were recorded as described by Study of Biological Diversity (UGICSBD), and Genoways and Williams (1979), including greatest University of Kansas Natural History Museum (KU). length of skull, zygomatic breadth, mastoid breadth, Each species account includes a list of specimens postorbital breadth, length of maxillary toothrow examined (sample size in parentheses) followed by (C-M3), and breadth across upper molars (M3-M3; see Region in Guyana, specific locality, and museum Table 2). catalogue numbers for specimens examined. Details of reproductive condition included testes (T) length by width in mm for males. For females we report presence or absence of an embryo with crown-rump length (CR) if present and note whether the individual was lactat- Family Emballonuridae ing. Details of capture, such as microhabitat, date, and time were recorded for each species. External mea- Centronycteris maximiliani (Fischer, 1829) surements (in mm) and body mass (in g) were recorded in the field as described in Nagorsen and Peterson Specimens examined (3). -Potaro-Sipa- (1980) and presented in Table 1, including total runi; Cleanvater Camp (KU 156014); Paka- length, length of tail, length of hindfoot, length of ear from notch, length of tragus,'length of forearm, and tau Falls (ROM 107081, 107082). body mass. Cranial measurements (in mm) for adult Two adult females were netted together specimens (complete ossification of phalanges and about 10 m above the ground in the forest

TABLE1. Range of six external measurements (in mm), including total length (TL), tail length (TAIL), hindfoot length (HFL), ear length (EAR), tragus length (TRAGUS), forearm length (FAL), and body mass (MASS, in g) of 10 new country records of bats for Guyana. Sample sizes differing from those reported under n are given in parentheses after the measurement or mass

Species n TL TAIL HFL EAR TRAGUS FAL MASS

Emballonuridae Centronycteris maximiliani 3 9 9 56-63(2) 17-23 6-7 14-17 4-6 44-46 5-6 Diclidurus albus 9 8 22 12 D. ingens 106 23 12 D. isabellus 83-92 15-24 12-14 86 20 11 Peropteryx leucoptera 56 5 8 59-61 9-1 1 7-9

Phyllostomidae Micronycteris brosseti 5 1 12 10 5 6 11 10 M. microtis 59 12 9 53-60 13-15 9-10 Tonatia carrikeri 77-95 9-16 15-17 74-88 15 16-17

Vespertilionidae Lasiurus atratus 115 5 2 10 Myotis riparius 78-85 35-39 7-10 80-83 33-38 8-9 182 B. K. Lim, M. D. Engstrom, R. M. Tim, R. P. Anderson, and L. C. Watson

TABLE 2. Range of six standard cranial measurements (in mm) of 10 new country records of bats for Guyana, including greatest length of skull (GLS), zygomatic breadth (ZB), mastoid breadth (MB), postorbital breadth (POB), length of maxillary toothrow (C-M'), and breadth across upper molars (M'-M') ~ - - Species n CLS ZB MB POB C-M' - Emballonuridae Cmtronycren's maximiiiani 29 P 14.7-14.9 8.7-8.8 7.2-7.3 3.G3.2 5.8-6.0 Dic1iduru.s olbur I 9 18.7 12.2 9.7 5.7 8.0 D. ingens 19 21.9 14.1 10.7 5.9 9.9

Id 21.3 13.5 10.8 5.9 9.6 D. isabelius 89 P 17.9-18.5 11.3-1 1.9 9.3-10.0 5.2-5.7 7.1-7.7 Peroprery leucoptera 19 - 9.5 8.0 3.3 6.2 Id 15.5 9.6 7.9 3.1 6.2 Phyllostomidac 17.6 8.5 8.1 3.8 6.6 17.5 8.3 8.0 3.8 6.7 18.2 9.1 8.4 4.0 6.9 18.3-18.7 8.6-9.1 8.3-8.5 3.84.2 6.8-7.1 23.9-25.4 11.0-11.5 11.4-12.2 3.54.0 79-86 25.6267 12.0-12.2 12.G13.4 3.9-4.1 8.7-8.8 Vespertilionidae 14.4 9.8 8.0 4.4 4.8 14.C-14.1 8.8-9.3 7.2-7.6 3.4-3.6 5.2-5.3 13.8-14.6 9.G9.5 7.2-7.6 3.4-3.7 5.2-5.5 - canopy at Pakatau Falls on 13 March 1997 at (Simmons and Handley, 1998). Previously, approximately 2100 h. A third adult female the genus was considered monotypic. How- was shot near the Burro Burro River while ever, based on cranial morphology including flying at about 1700 h in a slight depression a larger anterior upper premolar and smaller in semi-open forest near an old tree fall on 1 basisphenoid pits, C. centralis was consid- April 1997. None of these females was ered a distinct species found from southern pregnant or lactating. Mexico to Amazonian Peru (Simmons and Centronycteris maximiliani, as recently Handley, 1998).Their generic revisionincor- restricted by Simmons and Handley (1998), porated our three specimens from Guyana. is widely distributed but uncommon from southern Venezuela to southeastern Brazil. Diclidurus albus Wied-Neuwied, 1820 In the Guianan subregion, this species was known by one specimen from Venezuela Specimen examined (1). - Potaro-Sipa- (McCarthy and Ochoa, 1991), two from runi; Clearwater Camp (ROM 107435). Suriname (Husson, 1962; Williams et a/., One non-pregnant adult female was shot 1983), and one from French Guiana while flying high above the Burro Burro First records of 10 species of bats in Guyana 183

River early in the evening of 10 April 1997. Diclidurus isabellus (Thomas, 1920) One female Diclidurus ingens (see next Specimens examined (14). - Potaro-Si- account) was shot at the same time flying at paruni; Clearwater Camp (KU 154878- a similar height over the river, and four 154884; ROM 107389, 107390, 107436- female D. isabellus were also shot flying 107439; UGICSBD M161). lower over the river. Thirteen non-pregnant adult females and Diclidurus albus is rare but widely dis- one adult male (T = 2x1 mm) were shot tributed from Mexico to eastern Brazil while flying low over the Burro Burro River (Ceballos and Medellin, 1988). This species during the evenings of 4, 5, 6, and 10 April has been documented by only 24 specimens 1997. from four localities in Venezuela (Handley, This species was known previously by 1976; Ochoa G., 1995) and one specimen only the type specimen found in northwest- from Suriname (Ojasti and Linares, 1971). It em Brazil (Thomas, 1920) and 28 specimens has not been reported from French Gui- from three localities in Amazonas, Vene- ana. The external and cranial measure- zuela (Handley, 1976). There have been no ments for our specimen fit within the range reports from Suriname or French Guiana. reported by Cehallos and Medellin (1988) Our new records of D. isabellus for Guyana except for a marginally smaller mastoid increase the number of known specimens by breadth. another 50%, highlighting the collecting bias associated with some species of bats. We Diclidurus ingens Hemindez-Camacho. 1955 never caught this hat in mist nets or harp traps; however, collecting with a shotgun Specimens examined (2). - Demerara- indicates that they can he locally common, Mahaica; Georgetown (ROM 31981). Po- flying over relatively large open rivers. taro-Siparuni; Clearwater Camp (ROM Thomas (1920) described a new genus 107434). (Depanycteris) for D. isabellus because it Onenon-pregnant, adult female D. ingens was brownish in colour (not white as in other was shot while flying high above the Burro Diclidurus), had a more developed thumb, Burro River during the early evening on the relatively longer hindfoot compared to the 10th of April 1997. An adult male was tibia, morphologically distinct groove on the caught by Mr. Ram Singh after it flew into inner side of the tibia, and deeply emargi- a school classroom in Georgetown on 16 nated palate in the skull. Ojasti and Linares March 1961. (1971), however, suggested that the cranial Diclidurus ingens was known previously or external characteristics were not sufficient only by the holotype from southeastern to warrant generic separation of D. isabellus. Colombia (Hernandez-Camacho, 1955) and We follow Koopman (1993,1994) in placing fromvenezuelahy six specimens (Ojasti and this species in the genus Diclidurus. Linares, 1971; Handley, 1976). This species has been listed as occurring in Peru without Peropteiyx leucoptera Peters, 1867 reference to specimens (Pacheco et al., 1995) possibly because the type locality is on the Specimens examined (3). -Potaro-Sipa- border with northern Peru. It has not been runi; Iwokrama Field Station (ROM 107458). reported from Suriname or French Guiana. Upper Demerara-Berbice; Bada Creek, 18 The large size of this ghost bat sets it apart km NE Ituni (ROM 70688). Upper Takutu- from all other species of Diclidurus Upper Essequiho; Illia Wau River, 3 days (Ceballos and Medellin, 1988). below Kuitaro River mouth (ROM 41530). 184 B. K. Lim, M. D. Engstrom, R. M. Tirnm, R. P. Anderson, and L. C. Watson

An adult male (T = 3x2 mm) was netted and Peronymus as separate genera without at approximately 2000 h about 1 m above the further comment. ground along a trail in disturbed alluvial forest near Iwokrama Field Station on 12 April Family Phyllostomidae 1997. Additional specimens have been collected by Mr. Stan Brock who obtained an Micronycteris brosseti Simmons and adult female from the Illia Wau River in Voss, 1998 1967 and Mr. Baldwin Persaud who netted an adult male in Wallaba (Eperua sp.) bush Specimens examined (2). -Potaro-Sipa- near Bada Creek on 17 October 1973. * runi; Three Mile Camp (KU 155162; ROM This species is documented in northern 111469). South America east of the Andes by five One adult male (T = 2x1 mm) and one specimens from three localities in Peru non-pregnant adult female were caught in a (Thomas, 1924; Woodman et al., 1991), 34 mist net set in terra firme forest at Three specimens from three localities in the eastern Mile Camp on 24 March 1997. Amazon basin of Brazil (Sanborn, 1937; Micronycteris brosseti was recently Thomas, 1920), and one specimen from described from French Guiana with other Amazonian Colombia (Lemke et al., 1982). specimens known from eastern Peru and Three specimens were reported from two southeastern Brazil (Simmons and Voss, localities in Amazonas, Venezuela (Sanborn, 1998). Our two specimens from Guyana 1937; Ochoa G., 1984), as well as two speci- have pale grey ventral fur, short hairs on the mens from two localities in northeastern leading edge of the ears, and a calcar longer Suriname (Husson, 1962; Genoways et al., than the hindfoot. It is the smallest Micro- 1981), and twelve specimens from two nycteris presently known from Guyana. localities in northern French Guiana (Brosset We collected a single species of batfly, and Charles-Dominique, 1990; Simmons and Strebla alvarezi Wenzel, 1966 (Diptera: Voss, 1998). Streblidae) from M. brosseti. This poorly Miller (1907) elevated Peronymus, with known batfly seems to be associated with P. leucopterus as its only species, to the rank bats of the genus Micronycteris (Wenzel et of genus based on a less inflated rostrum, al., 1966), although this is the first report of connected ears, and white wings in compari- a parasite from M. brosseti. son with Peropteryx. Sanborn (1937) distin- guished Peronymus based on a relatively Micronycteris microtis Miller, 1898 broader rostrum and larger pterygoid pits at the anterior end of the basisphenoid pit. The Specimens examined (4). -Potaro-Sipa- taxon was synonymized with Peropteryx runi; Clearwater Camp (ROM 107354); by Cabrera (1958) without explanation. Three Mile Camp (ROM 107172); Kuru- Simmons and Voss (1998) followed this pukari (ROM 98159); Upper Takutu-Upper synonymy because the taxa share a unique Essequibo; Gunn' s Strip (ROM 106788). antebrachial wing sac morphology, and the One non-pregnant adult female was white wing membranes of P. leucoptera caught in a ground-level mist net in terra constitute a uniquely derived character. firme forest near Kurupukari on 20 October Based on hyoid morphology, Griffiths and 1990. Three adult males, all with T = 2x1 mm, Smith (199 1) suggested a sister-taxa relation- were caught in mist nets. One was netted ship based on two shared unique hyoid near the village of Gunn's Strip on 23 No- features; however, they retained Peropteryx vember 1996, another was caught in a First records of 10 species of bats in Guyana 185 canopy-level net at Three Mile Camp on 22 Tonatia carrikeri (J. A. Allen, 1910) March 1997, and the third was caught at Clearwater Camp on 3 April 1997. Specimens examined (9). -Potaro-Sipa- Micronycteris microtis was initially runi; Clearwater Camp (KU 155249-155251; described as a new species known from only ROM 107391, 107392); Three Mile Camp the holotype (Miller, 1898). Andersen (1906), (ROM 107190; UGICSBD M156); Iwo- however, questioned the validity of the krama Field Station (KU 155252; ROM condition of the ear as a diagnostic character 107461). and synonymized it under M. megalotis. A non-pregnant adult female and an adult With a larger sample size, Simmons (1996) male (T = 11x7 mm) were caught in a can- found the taxa occurring sympatrically in opy net set within terra firme forest at Three Colombia, Venezuela, French Guiana, and Mile Camp on 24 March 1997. Two non- Brazil, with the length of fur on the leading pregnant adult females were netted on edge of the ear distinguishing the two species 31 March and 2 April 1997, a lactating (M. microtis i4 mm). We tentatively follow female and a male (T = 4x2 mm) were netted this taxonomic arrangement but note that this on 6 April 1997, and a pregnant female (CR character is variable. Identifications are = 25 mm) was netted on 8 April 1997 at especially difficult given that, 'no consistent Clearwater Camp. Two non-pregnant adult differences in craniodental measurements or females were netted in alluvial forest near fur colour were found when large samples of the Iwokrarna Field Station on 12 April megalotis and microtis from many different 1997. localities were compared' (Simmons and As summarized by McCarthy et al. (1992), Voss, 1998: 76). We did not find the features T. carrikeri is found in northern South listed by Simmons and Voss (1998) for dist- America east of the Andes from northern inguishing sympatric species in French Bolivia, eastern Peru, Colombia, southern Guiana to be completely reliable in differen- Venezuela, Suriname, and northern Brazil. tiating these taxa in Guyana. More specifically, there are five reported As currently defined, M. microtis is localities from southern Venezuela, three distributed from southern Mexico to Amazo- for Suriname, and one specimen from nian Brazil (Simmons, 1996). It has been Roraima State in Brazil between southern reported from throughout southern Vene- Venezuela and Guyana. The specimen zuela (Handley, 1976) and French Guiana from Guyana reported by Swanepoel and (Brosset et al., 1996; Simmons, 1996). Genoways (1979) as T. carrikeri (ROM Micronycteris megalotis has been reported 67468) actually is T. schulzi (McCarthy from Suriname (Husson, 1962; Genoways et al., 1992). and Williams, 1986), but Brosset and Our nine specimens from Guyana and Charles-Dominique (1990) consider the the two reported from French Guiana specimens listed in Husson (1962) to be (Brosset et al., 1996; Simmons and Voss, referable to M. microtis based on the pub- 1998) indicate that T. carrikeri is dis- lished measurements. The distinction used tributed throughout the Guianan region. by Brosset and Charles-Dominique (1990) Tonatia carrikeri is the only species in of length of ear for M. megalotis averag- the genus with a pure white venter (McCar- ing 20 mm and 15 mm for M. microtis, thy et al., 1992), although older individ- however, is greater than the 22 mm versus uals of T. schulzi tend to have paler, al- 20 mm difference reported in Simmons most white, bellies (Simmons and Voss, (1996). 1998). 186 B. K. Lim, M. D. Engstrom, R. M. Timm, R. P. Anderson, and L. C. Watson

Family Vespertilionidae individuals were found roosting in a small crack and hollow in a branch of a 15 cm Lasiurus atratus Handley, 1996 diameter tree felled by local inhabitants near Specimens examined (1). -Potaro-Sipa- Surama Sawmill on 25 July 1994. All were mni; Three Mile Camp (ROM 107228). adults except for one juvenile male. Testes A non-pregnant adult female was caught measurements for six adult males ranged in a canopy net set over the road within terra from 7x5 to 10x8 mm, and two of five fe- fxme forest at Three Mile Camp on 27 males were pregnant with CR of 14 arid 20 March 1997. mm. One lactating adult female was caught This species was recently described based in a ground-level mist net set in the forest on the holotype from Suriname, 15 speci- beside standing water in an intermittent mens from two localities in southern Vene- stream near the Chodikar River on 30 Octo- zuela, and another reported specimen from ber 1996. Another lactating adult female was French Guiana (Handley, 1996). Our speci- caught in a ground-level mist net beside a men from Guyana has black wings, reddish creek about 7 km S of Gunn's Strip near the fur, and the range of measurements falls Essequibo River on 17 November 1996. An within those reported in Handley (1996) adult male (T = 8x6 mm) was caught in a except for greatest length of skull and fore- ground-level mist net set at Gunn's Strip on arm, which are both slightly larger. Lasiurus 21 November 1996. A pregnant female (CR egregius is larger still, with well developed = 11 mm) was caught in a mist net at lachrymal and supraorbital processes that are Clearwater Camp on 31 March 1997. lacking or poorly developed in L. atratus. As Myotis riparius is widely distributedfrom presently known, L. atratus is the only spe- Honduras to northern Uruguay (LaVal, cies of hat that is considered endemic to the 1973). It has been reported from throughout Guianan subregion. southern Venezuela (Handley, 1976) and French Guiana (Brosset and Charles- Myotis ripurius Handley, 1960 Dominique, 1990; Simmons and Voss, 1998) but not from Suriname although it is most Specimens examined (18). -Potaro -Si- surely also found there. The colour of the fur paruni; Clearwater Camp (ROM 107278); is quite variable, ranging from light grey- Surama Sawmill (ASNHC 9016-901 8,9020; brown to dark red-brown to black-brown, ROM 103133, 103136, 103138, 103150; making it difficult to distinguish some indi- UGICSBD M305; USNM F36598, F36604, viduals from M. rzigricuns in the field. F36605; these last three specimens have their Myotis albescens, however, has a distinctive field numbers reported as cross references frosting on the tips of its fur which distin- because they have not been officially depos- guishes it from the other two sympatrically- ited at the USNM); Kurupukari (BMNH occurring congeners in Guyana. Neverthe- 1997.42; Smith and Keny, 1996); 30 km less, the skull of M. riparius has a sagittal NE of Surama (ROM 98056). Upper crest, and the second upper premolar is Takutu-Upper Essequibo; Chodikar River minute, usually less than one-fourth the (ROM 106580); 7 km S of Gunn'~Strip height of the first premolar (LaVal, 1973). (ROM 106751); Gunn's Strip (ROM On average, M. ripurius is larger than M. 106773). nigricuns and M. albescens, with the An adult male (T = 8x5 mm) was caught zygomatic breadth (> 8.7 mm) and breadth in a ground-level mist net set in forest 30 km across upper canines (z 3.7 mm) consistently NE of Surama on 11 October 1990. Twelve larger. Additional characters useful in identi- First records of 10 species of bats in Guyana 187 fying neotropical Myotis are given in Timm Smith and Kerry (1996). Preliminary work and LaVal (1998). on the second Iwokrama Faunal Survey and on older collections alluded to by Parker et DISCUSSION al. (1993) suggests that additional species may occur in Iwokrama Forest and in Guy- Although the Royal Ontario Museum ana. (ROM) collected in the Iwokrama Forest Interestingly, the ten new species re- area in 1990, no published reports of mam- corded in Guyana are primarily insect feed- mals from this region existed until Smith and ing bats. Seven are considered aerial insecti- Kerry (1996) listed 36 species of bats as part vores, and three are gleaners. Micronycteris of a biodiversity survey by the Open Univer- microtis is probably relatively common in sity in 1992. They also included an addi- museum collections but is cryptic and diffi- tional nine species based on an unpublished cult to distinguish from M. megalotis. My- preliminary field report made by the ROM in otis riparius is also commonly captured, but 1990, bringing the total to 45 species of bats. the remaining eight species are not well Smith and Kerry (1996) extrapolated beyond represented in collections. Our success in their biodiversity survey of Iwokrama Forest using non-traditional collecting techniques and reported 96 species of bats from Guyana for bats such as large nets set high in based primarily on literature references. This the mid to upper canopy and shotguns total is reduced by one because, as discussed indicates the need to supplement ground- in Goodwin (1959), a single specimen from level nets to attain a more complete inven- New Amsterdam in Guyana that was origi- tory of species richness (Voss and Emmons, nally identified as Natalus stramineus, and 1996; Simmons and Voss, 1998). Likewise, cited in Jentinck (1893), Young (1896), estimates of relative abundances of species Beebe (19 l9), and Sanborn (1941), actually of bats can be greatly biased by survey is referable to Furipterus horrens, a previ- techniques. For example, prior to this study ously reported species from Guyana. Twelve D. isabellus was considered rare, but is of the 50 country records listed by Smith and actually locally common. As suggested by Kerry (1996) from outside Iwokrama Forest our study, the diversity of aerial insectivores were based on material that had not been in particular is underestimated by the tradi- recently reviewed (Parker et al., 1993). In tional technique of setting nets only 1-3 m addition to species listed by Smith and Kerry above the ground. (1996), as part of the description of Artibeus gnomus, Handley (1987) mentioned speci- GAZETTEER mens of this species from Guyana. Molossus pretiosus also has been reported from Guy- Bada Creek, 5"35'N, 58"08'W, 50 m a.s.1.; ana (Koopman, 1993, 1994). Our new re- Chodikar River, 1°22'N, 5g046'W, 240 m a.s.1.; cords bring the currently known bat fauna of Clearwater Camp, 4"44'N, 58"51'W, 90 m a.s.1.; Guyana to 107 species. In addition, there are Georgetown, 6"48'N, 58"10'W, at sea level; Gunn's Strip, 1°39'N, 58"38'W, 240 m a.s.1.; 29 new species records for Iwokrama Forest Gunn7sStrip, 7 km S, 1°35'N, 58"38'W, 240 m a.s.1.; bringing the total number of bats in this area Illia Wau River, 2"40'N, 5g040'W, -; to 74 species (B. K. Lim and M. D. Eng- Iwokrama Field Station, 4"40'N, 58"417W,70 m a.s.1.; strom, unpubl. data). The chiropteran diver- Kurupukari, 4"40'N, 5g040'W, 70 m a.s.1.; Pakatau Falls, 4"45'N, 59"01'W, 90 m a.s.1.; sity of Iwokrama Forest represents about Surama, 30 km NE, 4"20'N, 5g051'W, 70 m a.s.1.; 69% of the total bat diversity in Guyana, a Surama Sawmill, 4"06'N, 59"03'W, 80 m a.s.1.; larger proportion than the 43% reported by Three Mile Camp, 4"38'N, 5g043'W, 75 m a.s.1. 188 B. K. Lim, M. D. Engstrom, R. M. Timm, R. P. Anderson, and L. C. Watson

GENOWAYS,H. H., and S. L. WILLIAMS.1979. Re- cords of bats (Mammalia: Chiroptera) from We thank Daniel Allicock, Ron Allicock, Errol Suriname. Annals of Carnegie Museum, 48: McBirney, Dexter Torres, and Graham Watkins for 323-335. assisting us in the field. Collecting and export permits GENOWAYS,H. H., and S. L. WILLIAMS.1986. Results were facilitated by the staff at the Centre for the Study of the Alcoa Foundation-Suriname expeditions. of Biological Diversity, University of Guyana, Minis- XI. Bats of the genus Micronycteris (Mammalia: try of Agriculture, and Office of the President. The Chiroptera) in Suriname. Annals of Carnegie effort and dedication of Graham Watkins in coordinat- Museum, 55: 303-324. ing the faunal surveys for Iwokrama is gratefully GENOWAYS,H. H., S. L. WILLIAMS,and J. A. G'ROEN. acknowledged. We thank Matt Dean and Rupert 1981. Results of the Alcoa Foundation-Suriname Wenzel (The Field Museum) for providing the idehtifi- expeditions. V. Noteworthy records of Surina- cations of the batflies we collected. Constructive mese mammals. Annals of Carnegie Museum, 50: criticism was provided by two anonymous reviewers 3 19-332. and Wieslaw Bogdanowicz. Financial support was GOODWIN,G. G. 1959. Bats of the subgenus Natalus. provided by various sources including the Royal American Museum Novitates, 1977: 1-22.

Ontario Museum (ROM Foundation and Centre for GRIFFITHS,T. A., and A. L. SMITH.1991. Svstematics- J Biodiversity & Conservation Biology), the University of emballonuroid bats (Chiroptera: Emballonur- of Kansas Natural History Museum (E. Raymond Hall idae and Rhinopomatidae), based 01n hyoid mor- Fund), the U.S. National Science Foundation (Gradu- phology. Pp. 62-83, in Contributions to mamma- ate Fellowship to RPA), the Biological Diversity of logy in honor of Karl F. Koopman (T. A. GRIF- the Guianas Program at the Smithsonian Institution, FITHS and D. KLINGENER,eds.). Bi ulletin of the and The Academy of Natural Sciences of Philadelphia American Museum of Natural E[istory, 206: through the United Nations Development Program 1432. (UNDP). Collection management is ably provided by HANDLEY,C. O., JR. 1976. Mammals of the Smithso- the Mammalogy staffs at the ROM and KU. This is nian Venezuelan Project. Brigham Y oung Univer- contribution number 196 of the Centre for Biodiversity sity Science Bulletin, Biological S ieries, 20(5): and Conservation Biology at the Royal Ontario 1-91. Museum. HANDLEY,C. O., JR. 1987. New species of mammals from northern South America: fruit-eating bats, genus Artibeus Leach. Pp. 163-172, in Studies in Neotropical mammalogy: essays in hc onor of Philip ANDERSEN,K. 1906. On the bats of the genera Micro- Hershkovitz (B. D. PATTERSONand R. M. TIMM, nycteris and Glyphonycteris. Annals and Maga- eds.). Fieldiana, Zoology (N. S.), 39 : vii + 1-506. zine of Natural History, 7(18): 50-65. HANDLEY,C. O., JR. 1996. New species: of mammals

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Received 30 August, accepted 2 ~ecember1999