Occasional Papers Museum of Texas Tech University Number 249 18 January 2006 Bats of Antigua, Northern Lesser Antilles C. N. Scott Pedersen, Hugh H. Genoways, Matthew Morton, Vicki J. Swier, C. Peter A. Larsen, Kevel Lindsay, Rick A. Adams, and Jon D. Appino Introduction No comprehensive survey of the chiropteran was conducted, their work focused on roost surveys, fauna of the Antillean island of Antigua has been un¬ aiming to provide a resource for future work concerning dertaken in the past. Seven species of bats have been sites that would benefit from local conservation efforts reported in the literature from the island previously, and further study. but these have appeared in scattered reports. The first report of bats from Antigua was by Allen (1890) when The surveys conducted by parties led by Scott he reported a single specimen of Noctilio leporinus Pedersen in June 1998, July-August 2000, and June and two of Artibeus jamaicensis obtained during an 2003 provide the first significant results from mist-net¬ ornithological survey of Antillean islands. Subsequent ting bats in a variety of foraging habitats on Antigua. collections of bats on Antigua were incidental to herpe- Because the southwestern quadrant of the island was tological work or paleontological surveys of Antigua’s under-represented in previous survey efforts, the pri¬ limestone cave systems. We found 138 voucher speci¬ mary goal of the 2000 and 2003 surveys was to focus mens from Antigua scattered in eight museums from upon this forested region and possibly provide new previous collecting efforts. Before 2000, almost all species records for Antigua. The secondary goal of extant vouchers with locality data were collected from the survey was to provide comparative data for ongo¬ the vicinity of “Bats Cave,” situated on the southeast ing research into the impact of natural disasters on the corner of the island near the town of English Harbour bat population on the adjacent island of Montserrat in the Parish of St. Paul. The first formal survey of (Adams and Pedersen 1999; Pedersen 2001; Pedersen Antiguan bats was performed by Matthew Morton and et al., in prep.). Kevel Lindsay in 1994. Although some mist-netting Methods and Materials Study crretf.-Antigua is a small neotropical island measures a maximum of 21 km east to west and 16 km (279 km2) located in the northern Lesser Antilles in north to south. Boggy Peak (405 m) in the southwestern latitude 17°05'N and longitude 61°50'W. The island section of the island is the highest point. The mean 2 Occasional Papers, Museum of Texas Tech University annual temperature at St. John’s is 27.8°C, with the weather. Covered fiyways were netted wherever pos¬ highest monthly mean temperature in August (29.0°C) sible. At the end of each evening, bats were measured and the lowest in January (26. TC). The mean annual and examined (weight, forearm, reproductive status, rainfall for the island is 110 cm, with a dry season from tooth wear, presence of scars, and external parasites). January to April and a wet season from August to No¬ Together, the four mist-netting surveys included 78 vember. The island does lie in the hurricane zone and net-nights and yielded 157 captures of seven species received damage from Hurricane Jose and Hurricane of bat- Nocfilio leporinus, Monophyllusplethodon, Ar- Lenny in October 1999. Antigua is divided into three tibeus jamaicensis, Brachyphylla cavernarum, Nataius natural physiographic regions-a volcanic region in the stramineus, Tadarida brasiliensis, and Molossns mobs- southwest, a central plain, and a limestone region oc¬ sits. Sixty-four Artibew and nine Brachyphylla were cupying the northeastern third of the island ( Loveless wing-banded. During this time period, an additional 1960). Antigua was originally covered with low forest 56 bats, representing the seven species of bat reported or scrub, which was over time completely removed for from Antigua, were captured by hand in a variety of sugarcane plantations, pastures, or fuel. By the end of bat roosts, including Bats Cave, two rock overhangs the 18th century, nearly 142 km2 were under cultiva¬ on the west coast, and several buildings. tion, but by 1960, only 101 km2 were under cultivation leaving about 178 km2 covered by secondary types Limited radio-tracking of Brachyphylla was of vegetation that is maintained by clearing, shifting performed using 0.63 g BD-2 transmitters (Holohil cultivation, and the grazing of goats. Loveless (1960) Systems Ltd.) and TRX-48S Receivers (Wildlife Ma¬ suggested that there were originally a few areas of terials Inc.). closed-canopy. Evergreen Seasonal forest on Antigua. There are remnants of this vegetation in protected Voucher specimens. All voucher specimens mesic valleys on Boggy Peak, Rock Peak, Sugar Loaf from the 2000 and 2003 survey were deposited in the Mountain, and Wallings Hill. Antigua has no perma¬ research collections at the University of Nebraska nent rivers, but it does have more than 10 “seasonal” State Museum (UNSM) and the Museum of Texas watercourses, locally called ghauts. The most important Tech University (TTU). Other museum specimens of these is the semi-permanent Bendals River located were examined at the following institutions: AMNH, in the central plain (Loveless 1960). American Museum of Natural History, New York, NY; BMNH, British Museum (Natural History), London, Mist /7<?///>zg.-Mist-netting for bats was conducted UK; FMNH, Field Museum of Natural History, Chi¬ in a variety of habitats, including fruit plantations, cago, IL; KU, Natural History Museum, University of ponds, reservoirs, and access roads on four separate oc¬ Kansas, Lawrence, KS; MCZ, Museum of Comparative casions: 18 December-2 February 1994; 27 June 1998; Zoology, Harvard University, Cambridge, MA; MSU, 29 July - 6 August 2000; and 7-10 June 2003. The latter Museum of Michigan State University, East Lansing, two surveys were the most consistent in terms of netting MI; NMNH, National Museum of Natural History, effort with five to seven mist-nets being erected at each Washington, DC; UF, Florida Museum of Natural His¬ site and monitored for 4-6 h depending on activity and tory, University of Florida, Gainesville, FL. Results Capture Rates. Fruit bat captures per net-night neither the altitudinal gradient to generate rainfall, (BNN) range from 0.65 to 6.65 in the Neotropics (Find¬ nor heavily forested valleys whose native fruits might ley and Wilson 1983; Fenton et al. 1992; Pedersen, provide year-around support for additional species of unpublished data). However, phyllostomid captures on fruit bats as they do on the adjacent islands of Mont¬ Antigua were toward the low end of this scale at 1.45 serrat and Guadeloupe. Indeed, only fragmentary sec¬ BNN. It seems that Antigua’s flat terrain, abundant ondary growth forest covers the flanks of the Shekerley desert scrub, and anthropogenic disturbance provide Mountains and Boggy Peak (405 m) in the southwestern Pedersen et al.-Bats of Antigua, Northern Lesser Antilles 3 quarter of the island. These patches of forest were Animal Health.-Fruit bats on the adjacent island specifically targeted during the 2000 survey without of Montserrat have contended with the deposition of significantly different results (Fig. 1). volcanic ash on leaves, fruits, and flowers since the Figure 1. Map of Antigua indicating locations where bat research has been conducted. Numbers refer to locations shown in the figure. Closed circles represent mist-netting localities and active roost sites visited during the current study. Closed triangles are locations represented solely by museum voucher specimens from other studies. Open circles indicate inactive roosts visited. 1) Flat Top Hill: roost in rock overhang near Flat Top Hill. 2) Cades Bay (Goat Head): roost located in rock outcrop. 3) Cades Bay Plantation: fly ways located among mango trees at center of banana plantation at upper end of valley, 2.2 km ENE Urlings, 50 m in elevation. 4) Boggy Peak Road: fly way along densely covered access-road to radio installation on top of Boggy Peak, 3.0 km NE Urlings, 350 m. 5) Christian Valley Agricultural Station: access road located among banana trees, 2.1 km S Jennings, 50 m. 6) Christian Valley Agricultural Station: roost located in rock outcrop, northeastern slope of Christian Valley. 7) Green Castle Agricultural Station: fly way along water¬ works and unnamed stream leading towards Fiennes Well Water Station, Emanuel, 30 m. 8) Emmanuel: roost in derelict building southeast of the Green Castle Agricultural Station. 9) Claremont Plantation: flyways located among mango trees at center of banana plantation, 1.0 km NE Old Road, 20 m. 10) Wallings Reservoir: flyway along densely covered track leading up steep hillside south of reservoir and around reservoir margin, 1 km SSE John Hughes, 150 m. 11) Swetes Dam: fly way along densely covered access-road into Swetes Dam and across the top of the concrete dam itself, 1.5 km WNW Swetes, 50 m. 12) Garden in All Saints: garden at private residence in the town of All Saints, 50 m. 13) Bats Cave: large roost in limestone cave. 14) Shirley Heights: trail along crest of ridge. 15) Collins Reservoir: roost in derelict Mill. 16) Newfield: roost in derelict out-building behind Moravian Church. 17) Montpellier: natural limestone tunnel adjacent to the Montpellier sugar factory. 18) Santa Maria Hill. 19) English Harbour Town. 20) Mill Reef. 4 Occasional Papers, Museum of Texas Tech University Soufriere Hills volcano began to erupt in 1995. Al¬ fied guano were common. During the 1994, 1998, and though Antigua has received occasional wind-blown 2000 surveys, only two species of bat were observed ash from Montserrat during many of the larger eruptive in the cave-Brachyphylla cavernanun and Natalus events (1997-2003), none of the fruit bats captured on stramineus. In 2003, two species of bats also were Antigua in 2000 or 2003 exhibit any of the sub-lethal encountered, but they were Brachyphylla cavernarum pathologies associated with the ingestion of ash noted and Monophyllus plethodon. No Natal us stramineus on Montserrat (alopecia, dental attrition; Adams and were observed despite the excavation and exploration of Pedersen 1999; Pedersen 2001; Pedersen et al., in several side tunnels. This cave has a well-documented prep.). In Bats Cave on Antigua, random hand-captured history of utilization by both bats and humans (Nich¬ male, female, and juvenile Brachyphylla exhibited olson 1992). Indeed, this cave was outfitted with lights similar loads of external parasites (2-3 streblid flies, and developed as a tourist attraction in the 1970s, and Trie hob ins-, and numerous wing-mites), but exhibited the bat population was large enough that guano mining neither the extreme loads (15+ streblids), nor the ex¬ operations are reported back to 1897 (Nicholson 1992). tensive hair-loss noted on Brachyphylla on Montserrat Given the ease of access and the apparent tolerance of during this period (Pedersen et al., in prep.). Lactating the species to moderate disturbance. Bats Cave presents females often exhibit alopecia; however, Brachyphylla an extraordinary opportunity for further work on these on Montserrat have exhibited advanced alopecia since taxa. During their 1994 survey of Antigua and Barbuda, 1997-2000 whether they are lactating or not (Adams Morton and Lindsay found that 91.3% of all identified and Pedersen 1999; Pedersen, unpublished data). caves (11 on Antigua and 12 on Barbuda) were in use Although Brachyphylla on Antigua have encountered as bat roosts. This, combined with the fact that the larg¬ volcanic ash, lactating females were fully furred in est populations of bats on both islands were found in 2000 and 2003. This suggests that the levels of alopecia cave roosts, underlines the importance of these roosting noted in Brachyphylla on Montserrat are due to exces¬ habitats to bats. As such, Bats Cave is in great need of sive grooming and physiological stress rather than the protection, and will clearly benefit from conservation stress imposed by lactation alone. efforts by local authorities. Bats C<3v<?.~This large limestone cave is located Species Accounts.-The following species ac¬ on the southeast comer of the island near the town counts are drawn from census efforts by Morton of English Harbour. The surrounding habitat is des¬ and Lindsay in 1994, Pedersen and Adams in 1998, ert scrub, consisting mostly of Acacia. This cave is Pedersen, Appino, and Swier in 2000, and Pedersen, situated 1 km from the ocean, however, there is no Genoways, and Larsen in 2003. Detailed descriptions permanent source of fresh water within a 3 km radius of mist-netting habitats are provided in the legend ac¬ of the cave entrance. Beyond that distance, there are companying Figure 1. All length measurements are in numerous swimming pools located at resorts and pri¬ millimeters and mass is reported in grams. vate residences. The main elements of this cave consist of at least eight large domed chambers arranged in a Noctilio leporinus mastivus (Vahl 1797) somewhat linear fashion averaging 6-7 m in diameter extending nearly 60 m into the hillside. There are Specimen examined (1).-Parish Unknown: no numerous side chambers (possibly extensive given specific locality, 1 (FMNH). the geology) most of which were inaccessible to adult humans without excavation. The deepest portion of Specimens captured/released (4).St. Mary Par¬ the cave that was easily accessible by walking and ish: Cades Bay Plantation, 1; Flat Top Hill, I; Wallings crawling was approximately 15-20 m below ground Reservoir, 1. St. Paul Parish: Swetes Dam, 1 (Fig. 1). level. During the 2000 survey, the cave climate was 29.5° C with 81% relative humidity recorded in the Allen (1890) and Davis (1973) have reported this distal chamber. In 2000, guano deposits were between species previously from Antigua. Davis (1973), in his 15 and 45 cm in depth and supported a host of beetles, study of geographic variation in Noctilio leporinus, roaches, and flies. There was no evidence of ground assigned the one male that he examined from Antigua water, although small catchments of urine and lique¬ to the subspecies mastivus, with its type locality on Pedersen et al.-Bats of Antigua, Northern Lesser Antilles 5 St. Croix, Virgin Islands. The skull of our one specimen Monophyllus plethodon luciae Miller 1902 examined (FMNH 15083) is badly broken so that only the following measurements could be taken: length of Specimens examined(4).St Maty Parish: Chris¬ forearm, 84.6; postorbital constriction, 7.4; length of tian Valley, 2.1 km S Jennings, 32 m, 17°03'33"N, maxillary toothrow, 10.6; breadth across upper molars, 61°51 '44"W, 1 (TTU). St. Paul Parish: Bats Cave, 12.9. 1.5 km E English Harbour Town, 23 m, 17°00'52"N, 61°45'01"W, 2 (TTU). Parish Unknown: no specific Fishing bats were commonly observed along locality, 1 (MCZ). water courses on Antigua. Visual counts of Noctilio foraging over Swetes Dam and Wallings Reservoir in Specimens captnred/released (4).-St. Mary Par¬ 2000 demonstrate that mist-netting captures under¬ ish: Cades Bay Plantation, 1; Christian Valley Agri¬ estimate the Noctilio population at these sites by at cultural Stations, 1; Claremont Plantation, 1. St. Paul least a factor of three. The 13 million gallon Wallings Parish: Bats Cave, 1 (Fig. 1). Reservoir was constructed in 1890 and is surrounded by a mixed evergreen deciduous forest that was planted Schwartz and Jones (1967) first published on in 1915. Predominant tree species include locust (Hy- Monophyllus plethodon from Antigua based on four menaea courbarif), ironwood (Exostema caribaeum), specimens deposited in the British Museum (Natural mahoe (Daphnosis caribaea), black loblolly (Pisonia History) and Museum of Comparative Zoology, Har¬ fragrans), mango (Mangi/era indica), white cedar vard University. In their revision of the genus, they (Tabebuia pallida), mahogany (Swietenia mohogani), assigned these specimens to M. p. luciae, with its type and Spanish oak {Inga laurina). The Wallings Reservoir locality on St. Lucia. Later, Koopman (1968) also de¬ and Cades Bay Plantation mist-net captures in 1998 and scribed the specimen from Museum of Comparative 2000, respectively, were along densely wooded path¬ Zoology and provided selected measurements. Table 1 ways averaging a meter in width and several hundred presents measurements for three males from Antigua. meters away from the open water of the reservoir or the These measurements fall within the range of those of coastline. In all likelihood, these bats were commuting samples from Guadeloupe and Dominica (Baker et al. from unidentified roosts to their foraging habitat. Nei¬ 1978; Genoways et al. 2001), except for breadth across ther of the animals captured during July 2000 (female: the upper molars, which is broader in the specimens Cades Bay, male: Swetes Dam) evinced reproductive from Antigua. activity. In 2003, several fishing bats were observed foraging across the Wallings Reservoir but none were Average length of forearm and body mass for captured and one bounced off a mist net set across the three males that were captured and released was the south end of the reservoir. The forearm lengths 40.6 (40.0-42.0) and 16.7 (16.0-17.6), respectively. and body masses for the two males (Swetes Dam and Considering the number of nights spent mist-netting Wallings Reservoir) were 89.0 and 86.0, and 71.1 and in fruit orchards across a range of elevations (20-250 68.5, respectively, whereas the length of forearm and m), we were surprised at the relatively small number of body mass for the female from Cades Bay was 87.0 Monophyllus netted on Antigua in 2000 (4% of phyl- and 58.5. lostomid bat captures) in comparison to collections at similar sites on neighboring Montserrat in July 2000 In 1994, a non-reproductive male was extracted (14% of phyllostomid bat captures). from a shallow fissure located under a rock overhang near Flat Top Hill during daylight hours by Morton In 2003, three males were captured-two in Bats and Lindsay. They observed Noctilio roosting with Cave (June 8) and one netted in fruit orchards in Chris¬ the body pressed flat against rock surfaces. These bats tian Valley (June 9). The Monophyllus taken in Bats had been seen returning to Flat Top Hill to roost after Cave were found in a small side tunnel that opened dark. During a search of the adjacent cliff face, they in the floor of the right-hand chamber where no other observed Noctilio in large holes and fissures extending species were found. This small descending tunnel was upward into rock overhangs, although Noctilio was approximately 1 m wide and 1.5 m high, at least 30 never observed wedged into narrow cracks. m long, and may have had a separate opening to the 6 Occasional Papers, Museum of Texas Tech University outside as evidenced by a slight breeze in that tunnel. Brachyphylla cavernarum numbering well over 20,000 At least 20 individual Monophyllus were observed animals in Bats Cave (independent estimates by three in this small space. Three captured males had testes observers). In contrast to the Brachyphylla maternity lengths of 5,4, and 3 and weighed 15.5, 13.6, and 15.4, colony on the neighboring island of Montserrat, the respectively. Brachyphylla colony on Antigua was not separated into male/female caves, nor was there any obvious Brachyphylla cavernarum cavernarum evidence of male-female territoriality. Animals were Gray 1834 observed in clusters ranging in size from 10 to 200 individuals. The clusters of animals occupied numer¬ Specimens examined (114).-St. Paul Parish: ous pockets in the domed roof of each chamber of Bats Cave, 1.5 km E English Harbour Town, 23 m, the cave. Neonates and juvenile animals appeared to 17°00/52//N, 6r45'01"W, 20 (16 TTU, 4 UNSM); be randomly located throughout the cave, giving no Bats Cave, 2 mi. E Falmouth, 5 (2 FMNH, 3 UF); near indication of a “creche.” Numerous mid-air collisions Bats Cave, English Harbor [= Harbour], 12 (NMNH); between individual Brachyphylla were observed, sev¬ 1 mi. E English Harbor [=Harbour], 3 (1 KU, 2 MSU). eral resulting in the grounding of both animals. Due Parish Unknown: no specific locality, 74 (1 BMNH, to the apparent difficulty in taking flight from the soft 73 NMNH). substrate (guano), most grounded bats climbed up onto an adjacent rock surface before taking off. Of note, the Specimens captured/released (105).-St. Mary exposed upper surfaces of most rocks in lower half of Parish: Boggy Peak Road, 1; Cades Bay Plantation, 6; the cave were highly burnished, no doubt in response Christian Valley Agriculture Station, 1; Wallings Res¬ to animal contact. ervoir, 2. St. Paul Parish: Bats Cave, 95 (Fig. 1). Bats Cave was visited on 8 June 2003, when the Miller (1913a; see also Koopman 1968) was majority of the females in the colony were carrying the first author to report Brachyphylla cavernarum pups. The females were located in the deeper, hotter, from Antigua. Swanepoel and Genoways (1978) more humid portions of the cave. Many of the males examined 20 specimens from the island as part of were congregated near the entrance and in the outer¬ their systematic review of the genus. Swanepoel and most chambers where the temperature was considerably Genoways (1978) assigned the material from Antigua lower and ambient light made the bats clearly visible, to the nominate subspecies, which has a type locality Five of the six females obtained in 2003 were lactat- of St, Vincent. Table 1 presents the length of forearm ing and were collected along with their infants. The and seven cranial measurements for 12 males and sixth female was non-reproductive. The testes length 9 females from Antigua. The only measurement in of six adult males collected at this time averaged 6.3 which there was a significant difference between the (4-9). The length of forearm and weight of three male sexes was mastoid breadth, with males being larger and one female infant were 56.6, 51.6,51.8, and 52.2 (P<0.05). In general, males were larger in all other and 25.6, 21.8, 20.6, and 20.7, respectively. A male cranial measurements except postorbital constriction captured on 25 June 1958, had testes that were 5.0 in which the sexes were similar. Females had longer long. Of 10 females collected and preserved from Bats forearms. The measurements of our sample from An¬ Cave on 29 April 1959, nine were pregnant. Some of tigua match closely those samples of B. c. cavernarum the embryos had been removed for other studies, but presented by Swanepoel and Genoways (1978). the crown-rump length of remaining four embryos was 34, 36, 37, and 39. All 21 females collected on 1 May Sex ratio data for Brachyphylla hand-captured 1958 were pregnant, with one of the embryos measur¬ in 2000 in Bats Cave suggest near male-female par¬ ing 37 in crown-rump length. None of the nine females ity (48 males: 44 females). This male-female balance taken on 2 and 7 August 1903, the 3 females taken on was mirrored by a 5:4 male-female capture ratio while 7 November 1903, or the 5 females from 20 November mist netting in several foraging habitats located many 1903 were pregnant. These combined reproductive kilometers away from the cave. On 3 August 2000, data suggest that a monestrus reproductive pattern we observed a mixed male/female maternity colony of with synchronous breeding (Wilson 1979) occurs in Brachyphylla on Antigua. Pedersen et al.-Bats of Antigua, Northern Lesser Antilles 7 Average length of forearm and body mass (1994 male captured on 30 December 1889, and an adult fe¬ and 2000 capture/release) for 12 adult males was 66.2 male captured on 23 April 1890. Genoways et al. (2001) (64.6-69.0) and 49.5 (41.8-56.0), and the same for six reviewed Antillean populations of the Jamaican fruit adult females was 66.2 (64.0-68.0) and 46.4 (41.1- bat based upon morphometries and presence/absence of 49.4), respectively. Five lactating females captured in M3/m3 and Phillips et al. (1989) and Pumo et al. (1996) early June 2003 weighed an average of42.7 (38.6-46.3), presented genetic data for these populations. These whereas a non-reproductive adult female weighed 36.5. studies support the use of the subspecific name A. j. Six adult males taken at this time weighed an average jamaicensis for Antiguan populations. Table 1 presents of 43.4 (39.0-48.5). length of forearm and seven cranial measurements for four male and five female A. jamaicensis from Antigua. Morton and Lindsay recorded roost emergence The sexes differed significantly only in postorbital behavior for Brachvphylla from Bats Cave in February constriction in which females were larger on average. 1994. They observed that a small number of bats began Females were larger on average than males in all other to exit the cave to forage around 1845 h, whereas the measurements as well except length of the maxillary majority of the colony waited until after dark (1915 h) toothrow in which the sexes had the same mean. to exit the cave. The emergence continued sporadically through 1945 h. Eighteen of the 29 males (62%) captured and released in July 2000 exhibited scrotal testes. Of the 35 Radio-tracking data for two male Brachvphylla females captured and released in July 2000, five were indicate that these robust bats travel nightly up to 22 pregnant and 11 were lactating or post-lactation (45% km away from Bats Cave to forage in the moist, higher of the females were reproductively active). Six adult altitude valleys and farm lands on the southwestern males collected between 7-9 June 2003, had testes that side of the island. In addition, one of the wing-banded averaged 9 (6-11) in length. Of six females taken during Brachyphylla was recaptured at 14.3 and 15.4 km from this same time period, three evinced no reproductive the roost on consecutive evenings (minimum-commut¬ activity, two were lactating, and one carried an embryo ing distance of 60 km over two nights). that was 20 mm in crown-rump length. Two females taken at Santa Maria Hill on 14-15 August 1980 were Artibeus jamaicensis jamaicemis lactating. Leach 1821 For individuals that were captured and released, Specimens examined (28).-St. John Parish: the average length of forearm and body mass for 12 Santa Maria Hill, 2+ mi. NE St. Johns, 4 (NMNH). adult males were 59.5 (57.0-63.0) and 40.9 (36.3-50.6), St. Mary Parish: Christian Valley, 2.1 km S Jennings, respectively, and the same for 26 adult non-pregnant 32 m, 17°03'33"N, 61°51'44"W, 6 (TTU); Wallings females were 60.4 (57.0-62.0) and 42.5 (34.7-50.4), Reservoir, Wallings, 166 m, 17°02'04"N, 61°49'30"W, respectively. The weights of three non-pregnant females 6 (TTU). St. Paul Parish: English Harbor [= Harbour], taken in early June 2003 were 34.7, 36.9, and 36.9, 10(1 MCZ, 9 NMNH). Parish Unknown: no specific whereas two lactating females weighed 39.7 and 39.9. locality, 2(1 FMNH, 1 MCZ). Six adult males, also taken in early June 2003, weighed an average of 38.9 (35.5-43.2). Specimens captured/released (80).-St. John Parish: Green Castle Agricultural Station, 7. St. Mary Although W. R. Forrest recorded in his field Parish: Boggy Peak Road, 9; Cades Bay Plantation, 6; notes that Artibeus was present in “Bats Cave” in 1918, Christian Valley Agriculture Station, 10; Claremont Artibeus was not observed there in 1994, 1998, 2000, Plantation, 11; Wallings Reservoir, 12. St. Paul Parish: or 2003. However, Artibeus was commonly observed All Saints, 4; Swetes Dam, 16. St. Philip Parish: Col¬ in abandoned buildings and was collected at all mist- lins, 1; Montpellier, 4 (Fig. 1; Table 1). netting localities in 1994, 1998,2000, and 2003. These animals were observed hanging pendant from foliage J. A. Allen (1890, see also G. M. Allen 1911) first and from the ceilings/rafters of unused buildings, either reported this species from Antigua based on an adult individually or in very tight clusters. 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In direct contrast to the raucous nature Cave, while two others were hand-collected in a large of BrachyphyUa roosts, Artibens roosts typically remain rock crevice between very large boulders in Christian quiet unless the animals are disturbed by attempts to Valley in 1994 (1600 h) (Fig. 1). In 2000, numerous capture them. Natalus (n ~= 90+) were observed in Bats Cave, but no young were observed. As is typical of the species, Of the eight individuals examined, seven were each bat hung fully pendant a short distance (15-20 missing both upper third molars and possessed both cm) from neighboring bats across the ceiling of Bats lower third molars. The eighth individual was missing Cave. In 2000, Natalus was again observed to utilize both third upper molars as well as the lower left third height restricted portions of the cave ceiling along large molar. The lower right molar was minute. On Antigua, fissures that diverge laterally from the main chamber. heavy tooth wear was noted only on a single female The apparent preference for these smaller side passages Artibeus in 2000 (1% of captures) that was clearly a along the middle and at the back of the cave may be very old animal. due to a variety of reasons including: (1) the slightly higher relative humidity in these microclimates (85% Natalus stramineus stramineus Gray 1834 humidity), and (2) a desire to avoid the much larger BrachyphyUa. Indeed, we witnessed a mid-air collision Specimens examined (8).-St. Paul Parish: Bats between Natalus and BrachyphyUa that fractured the Cave, 1.5 km English Harbour Town, 23 m, 17°00'52"N, right humerus of the smaller bat (UNSM 27900). When 6r45'01"W, 1 (UNSM); English Harbor [= Harbour], funnel-eared bats began emerging around 1840 h, they 2 (MCZ). St. Philip Parish: Montpelier Cave, 1.2 km were flying low to the ground, but did not appear to have E, 0.2 km S, 78 m, 17°02'36"N, 61°425'2"W, 3 (TTU). made use of the main exit hole. The facultative use of Parish Unknown: no specific locality, 2 (MCZ). secondary exits may avoid collisions with BrachyphyUa that also began to exit around this time. No Natalus Specimens captured/released (9).-St. Mary Par¬ were observed in Bats Cave on 8 June 2003. ish: Christian Valley, 2. St. Paul Parish: Bats Cave, 7. During the 1998 and 2003 surveys, a limestone Goodwin (1959), in his revision of members of cave 600 m east of the disused Montpellier sugar fac¬ the subgenus Natalus, restricted the type locality of tory east of St. Philips was visited. This tunnel ran for Natalus stramineus to Antigua rather than Lagoa Santa, nearly 120 m with a relatively constant diameter of 1.5 Minas Gerais, Brazil, as was done by earlier authors. to 2.0 m. At least five Natalus were observed in this With this decision, which later was reconfirmed by cave in 1998 and 12 to 15 animals were observed in Handley and Gardner (1990), the nominate subspe¬ 2003. The northeastern half of Antigua is dominated by cies N. s. stramineus becomes the appropriate name to xeric habitats like those surrounding the Montpellier apply to the population on Antigua. Goodwin (1959) Cave and Bats Cave. These caves are very important gives measurements for two males from Antigua in the from a management perspective as they provide criti¬ British Museum (Natural History) as follows: length of cal humid microenvironments that Natalus must have forearm, 38.0,38.8; greatest length of skull, 16.6, 16.6: across this xeric part of the island. condylobasal length, 15.5, 15.7; zygomatic breadth, 8.7, 8.8; breadth of braincase, 8.1, 8.0; interorbital Length of forearm and body mass for two fe¬ breadth, 3.4,3.5; breadth across upper molars, 5.9, 5.9; males captured/released in 1994 and 2000 were 39.0 length of maxillary toothrow, 7.4, 7.5. Table 1 presents and 40.0 and 5.2 and 6.5, respectively. A female and the length of forearm and seven cranial measurements two males taken on 9 June 2003, weighed 4.4, 4.8, for three males and three females from Antigua. The and 5.0, respectively. The female evinced no gross range of measurements of these topotypic individuals reproductive activity and the males had testes lengths fall within the range of a larger sample from Dominica of 1.0 and 2.0. (Genoways et al. 2001).