Bats of Antigua, northern Lesser Antilles

Occasional Papers

Museum of Texas Tech University

Number 249 18 January 2006

Bats of Antigua, Northern Lesser Antilles

Scott C. Pedersen, Hugh H. Genoways, Matthew N. Morton, Vicki J. Swier,
Peter A. Larsen, Kevel C. Lindsay, Rick A. Adams, and Jon D. Appino

Introduction

No comprehensive survey of the chiropteran
fauna of the Antillean island of Antigua has been un¬
dertaken in the past. Seven species of bats have been
reported in the literature from the island previously,
but these have appeared in scattered reports. The first
report of bats from Antigua was by Allen (1890) when
he reported a single specimen of Noctilio leporinus
and two of Artibeus jamaicensis obtained during an
ornithological survey of Antillean islands. Subsequent
collections of bats on Antigua were incidental to herpe-
tological work or paleontological surveys of Antigua’s
limestone cave systems. We found 138 voucher speci¬
mens from Antigua scattered in eight museums from
previous collecting efforts. Before 2000, almost all
extant vouchers with locality data were collected from
the vicinity of “Bats Cave,” situated on the southeast
corner of the island near the town of English Harbour
in the Parish of St. Paul. The first formal survey of
Antiguan bats was performed by Matthew Morton and
Kevel Lindsay in 1994. Although some mist-netting

was conducted, their work focused on roost surveys,
aiming to provide a resource for future work concerning
sites that would benefit from local conservation efforts
and further study.

The surveys conducted by parties led by Scott
Pedersen in June 1998, July-August 2000, and June
2003 provide the first significant results from mist-net¬
ting bats in a variety of foraging habitats on Antigua.
Because the southwestern quadrant of the island was
under-represented in previous survey efforts, the pri¬
mary goal of the 2000 and 2003 surveys was to focus
upon this forested region and possibly provide new
species records for Antigua. The secondary goal of
the survey was to provide comparative data for ongo¬
ing research into the impact of natural disasters on the
bat population on the adjacent island of Montserrat
(Adams and Pedersen 1999; Pedersen 2001; Pedersen
et al., in prep.).

Methods and Materials

Study crretf.-Antigua is a small neotropical island
(279 km 2 ) located in the northern Lesser Antilles in
latitude 17°05'N and longitude 61°50'W. The island

measures a maximum of 21 km east to west and 16 km
north to south. Boggy Peak (405 m) in the southwestern
section of the island is the highest point. The mean

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Occasional Papers, Museum of Texas Tech University

annual temperature at St. John’s is 27.8°C, with the
highest monthly mean temperature in August (29.0°C)
and the lowest in January (26. TC). The mean annual
rainfall for the island is 110 cm, with a dry season from
January to April and a wet season from August to No¬
vember. The island does lie in the hurricane zone and
received damage from Hurricane Jose and Hurricane
Lenny in October 1999. Antigua is divided into three
natural physiographic regions-a volcanic region in the
southwest, a central plain, and a limestone region oc¬
cupying the northeastern third of the island ( Loveless
1960). Antigua was originally covered with low forest
or scrub, which was over time completely removed for
sugarcane plantations, pastures, or fuel. By the end of
the 18th century, nearly 142 km 2 were under cultiva¬
tion, but by 1960, only 101 km 2 were under cultivation
leaving about 178 km 2 covered by secondary types
of vegetation that is maintained by clearing, shifting
cultivation, and the grazing of goats. Loveless (1960)
suggested that there were originally a few areas of
closed-canopy. Evergreen Seasonal forest on Antigua.
There are remnants of this vegetation in protected
mesic valleys on Boggy Peak, Rock Peak, Sugar Loaf
Mountain, and Wallings Hill. Antigua has no perma¬
nent rivers, but it does have more than 10 “seasonal”
watercourses, locally called ghauts. The most important
of these is the semi-permanent Bendals River located
in the central plain (Loveless 1960).

Mist /7<?///>zg.-Mist-netting for bats was conducted
in a variety of habitats, including fruit plantations,
ponds, reservoirs, and access roads on four separate oc¬
casions: 18 December-2 February 1994; 27 June 1998;
29 July - 6 August 2000; and 7-10 June 2003. The latter
two surveys were the most consistent in terms of netting
effort with five to seven mist-nets being erected at each
site and monitored for 4-6 h depending on activity and

weather. Covered fiyways were netted wherever pos¬
sible. At the end of each evening, bats were measured
and examined (weight, forearm, reproductive status,
tooth wear, presence of scars, and external parasites).
Together, the four mist-netting surveys included 78
net-nights and yielded 157 captures of seven species
of bat- Nocfilio leporinus, Monophyllusplethodon , Ar-
tibeus jamaicensis, Brachyphylla cavernarum, Nataius
stramineus , Tadarida brasiliensis, and Molossns mobs-
sits. Sixty-four Artibew and nine Brachyphylla were
wing-banded. During this time period, an additional
56 bats, representing the seven species of bat reported
from Antigua, were captured by hand in a variety of
bat roosts, including Bats Cave, two rock overhangs
on the west coast, and several buildings.

Limited radio-tracking of Brachyphylla was
performed using 0.63 g BD-2 transmitters (Holohil
Systems Ltd.) and TRX-48S Receivers (Wildlife Ma¬
terials Inc.).

Voucher specimens. All voucher specimens
from the 2000 and 2003 survey were deposited in the
research collections at the University of Nebraska
State Museum (UNSM) and the Museum of Texas
Tech University (TTU). Other museum specimens
were examined at the following institutions: AMNH,
American Museum of Natural History, New York, NY;
BMNH, British Museum (Natural History), London,
UK; FMNH, Field Museum of Natural History, Chi¬
cago, IL; KU, Natural History Museum, University of
Kansas, Lawrence, KS; MCZ, Museum of Comparative
Zoology, Harvard University, Cambridge, MA; MSU,
Museum of Michigan State University, East Lansing,
MI; NMNH, National Museum of Natural History,
Washington, DC; UF, Florida Museum of Natural His¬
tory, University of Florida, Gainesville, FL.

Results

Capture Rates. Fruit bat captures per net-night
(BNN) range from 0.65 to 6.65 in the Neotropics (Find¬
ley and Wilson 1983; Fenton et al. 1992; Pedersen,
unpublished data). However, phyllostomid captures on
Antigua were toward the low end of this scale at 1.45
BNN. It seems that Antigua’s flat terrain, abundant
desert scrub, and anthropogenic disturbance provide

neither the altitudinal gradient to generate rainfall,
nor heavily forested valleys whose native fruits might
provide year-around support for additional species of
fruit bats as they do on the adjacent islands of Mont¬
serrat and Guadeloupe. Indeed, only fragmentary sec¬
ondary growth forest covers the flanks of the Shekerley
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
specifically targeted during the 2000 survey without
significantly different results (Fig. 1).

Animal Health .-Fruit bats on the adjacent island
of Montserrat have contended with the deposition of
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.

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Occasional Papers, Museum of Texas Tech University

Soufriere Hills volcano began to erupt in 1995. Al¬
though Antigua has received occasional wind-blown
ash from Montserrat during many of the larger eruptive
events (1997-2003), none of the fruit bats captured on
Antigua in 2000 or 2003 exhibit any of the sub-lethal
pathologies associated with the ingestion of ash noted
on Montserrat (alopecia, dental attrition; Adams and
Pedersen 1999; Pedersen 2001; Pedersen et al., in
prep.). In Bats Cave on Antigua, random hand-captured
male, female, and juvenile Brachyphylla exhibited
similar loads of external parasites (2-3 streblid flies,
Trie hob ins-, and numerous wing-mites), but exhibited
neither the extreme loads (15+ streblids), nor the ex¬
tensive hair-loss noted on Brachyphylla on Montserrat
during this period (Pedersen et al., in prep.). Lactating
females often exhibit alopecia; however, Brachyphylla
on Montserrat have exhibited advanced alopecia since
1997-2000 whether they are lactating or not (Adams
and Pedersen 1999; Pedersen, unpublished data).
Although Brachyphylla on Antigua have encountered
volcanic ash, lactating females were fully furred in
2000 and 2003. This suggests that the levels of alopecia
noted in Brachyphylla on Montserrat are due to exces¬
sive grooming and physiological stress rather than the
stress imposed by lactation alone.

Bats C<3v<?.~This large limestone cave is located
on the southeast comer of the island near the town
of English Harbour. The surrounding habitat is des¬
ert scrub, consisting mostly of Acacia. This cave is
situated 1 km from the ocean, however, there is no
permanent source of fresh water within a 3 km radius
of the cave entrance. Beyond that distance, there are
numerous swimming pools located at resorts and pri¬
vate residences. The main elements of this cave consist
of at least eight large domed chambers arranged in a
somewhat linear fashion averaging 6-7 m in diameter
extending nearly 60 m into the hillside. There are
numerous side chambers (possibly extensive given
the geology) most of which were inaccessible to adult
humans without excavation. The deepest portion of
the cave that was easily accessible by walking and
crawling was approximately 15-20 m below ground
level. During the 2000 survey, the cave climate was
29.5° C with 81% relative humidity recorded in the
distal chamber. In 2000, guano deposits were between
15 and 45 cm in depth and supported a host of beetles,
roaches, and flies. There was no evidence of ground
water, although small catchments of urine and lique¬

fied guano were common. During the 1994, 1998, and
2000 surveys, only two species of bat were observed
in the cave-Brachyphylla cavernanun and Natalus
stramineus. In 2003, two species of bats also were
encountered, but they were Brachyphylla cavernarum
and Monophyllus plethodon. No Natal us stramineus
were observed despite the excavation and exploration of
several side tunnels. This cave has a well-documented
history of utilization by both bats and humans (Nich¬
olson 1992). Indeed, this cave was outfitted with lights
and developed as a tourist attraction in the 1970s, and
the bat population was large enough that guano mining
operations are reported back to 1897 (Nicholson 1992).
Given the ease of access and the apparent tolerance of
the species to moderate disturbance. Bats Cave presents
an extraordinary opportunity for further work on these
taxa. During their 1994 survey of Antigua and Barbuda,
Morton and Lindsay found that 91.3% of all identified
caves (11 on Antigua and 12 on Barbuda) were in use
as bat roosts. This, combined with the fact that the larg¬
est populations of bats on both islands were found in
cave roosts, underlines the importance of these roosting
habitats to bats. As such, Bats Cave is in great need of
protection, and will clearly benefit from conservation
efforts by local authorities.

Species Accounts.- The following species ac¬
counts are drawn from census efforts by Morton
and Lindsay in 1994, Pedersen and Adams in 1998,
Pedersen, Appino, and Swier in 2000, and Pedersen,
Genoways, and Larsen in 2003. Detailed descriptions
of mist-netting habitats are provided in the legend ac¬
companying Figure 1. All length measurements are in
millimeters and mass is reported in grams.

Noctilio leporinus mastivus (Vahl 1797)

Specimen examined (1).-Parish Unknown: no
specific locality, 1 (FMNH).

Specimens captured/released (4).St. Mary Par¬
ish: Cades Bay Plantation, 1; Flat Top Hill, I; Wallings
Reservoir, 1. St. Paul Parish: Swetes Dam, 1 (Fig. 1).

Allen (1890) and Davis (1973) have reported this
species previously from Antigua. Davis (1973), in his
study of geographic variation in Noctilio leporinus,
assigned the one male that he examined from Antigua
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
examined (FMNH 15083) is badly broken so that only
the following measurements could be taken: length of
forearm, 84.6; postorbital constriction, 7.4; length of
maxillary toothrow, 10.6; breadth across upper molars,
12.9.

Fishing bats were commonly observed along
water courses on Antigua. Visual counts of Noctilio
foraging over Swetes Dam and Wallings Reservoir in
2000 demonstrate that mist-netting captures under¬
estimate the Noctilio population at these sites by at
least a factor of three. The 13 million gallon Wallings
Reservoir was constructed in 1890 and is surrounded
by a mixed evergreen deciduous forest that was planted
in 1915. Predominant tree species include locust ( Hy -
menaea courbarif ), ironwood ( Exostema caribaeum ),
mahoe ( Daphnosis caribaea ), black loblolly (Pisonia
fragrans), mango (Mangi/era indica ), white cedar
(Tabebuia pallida ), mahogany ( Swietenia mohogani ),
and Spanish oak {Inga laurina ). The Wallings Reservoir
and Cades Bay Plantation mist-net captures in 1998 and
2000, respectively, were along densely wooded path¬
ways averaging a meter in width and several hundred
meters away from the open water of the reservoir or the
coastline. In all likelihood, these bats were commuting
from unidentified roosts to their foraging habitat. Nei¬
ther of the animals captured during July 2000 (female:
Cades Bay, male: Swetes Dam) evinced reproductive
activity. In 2003, several fishing bats were observed
foraging across the Wallings Reservoir but none were
captured and one bounced off a mist net set across
the south end of the reservoir. The forearm lengths
and body masses for the two males (Swetes Dam and
Wallings Reservoir) were 89.0 and 86.0, and 71.1 and
68.5, respectively, whereas the length of forearm and
body mass for the female from Cades Bay was 87.0
and 58.5.

In 1994, a non-reproductive male was extracted
from a shallow fissure located under a rock overhang
near Flat Top Hill during daylight hours by Morton
and Lindsay. They observed Noctilio roosting with
the body pressed flat against rock surfaces. These bats
had been seen returning to Flat Top Hill to roost after
dark. During a search of the adjacent cliff face, they
observed Noctilio in large holes and fissures extending
upward into rock overhangs, although Noctilio was
never observed wedged into narrow cracks.

Monophyllus plethodon luciae Miller 1902

Specimens examined(4).St Maty Parish: Chris¬
tian Valley, 2.1 km S Jennings, 32 m, 17°03'33"N,
61°51 '44"W, 1 (TTU). St. Paul Parish: Bats Cave,
1.5 km E English Harbour Town, 23 m, 17°00'52"N,
61°45'01"W, 2 (TTU). Parish Unknown: no specific
locality, 1 (MCZ).

Specimens captnred/released (4).-St. Mary Par¬
ish: Cades Bay Plantation, 1; Christian Valley Agri¬
cultural Stations, 1; Claremont Plantation, 1. St. Paul
Parish: Bats Cave, 1 (Fig. 1).

Schwartz and Jones (1967) first published on
Monophyllus plethodon from Antigua based on four
specimens deposited in the British Museum (Natural
History) and Museum of Comparative Zoology, Har¬
vard University. In their revision of the genus, they
assigned these specimens to M. p. luciae , with its type
locality on St. Lucia. Later, Koopman (1968) also de¬
scribed the specimen from Museum of Comparative
Zoology and provided selected measurements. Table 1
presents measurements for three males from Antigua.
These measurements fall within the range of those of
samples from Guadeloupe and Dominica (Baker et al.
1978; Genoways et al. 2001), except for breadth across
the upper molars, which is broader in the specimens
from Antigua.

Average length of forearm and body mass for
the three males that were captured and released was
40.6 (40.0-42.0) and 16.7 (16.0-17.6), respectively.
Considering the number of nights spent mist-netting
in fruit orchards across a range of elevations (20-250
m), we were surprised at the relatively small number of
Monophyllus netted on Antigua in 2000 (4% of phyl-
lostomid bat captures) in comparison to collections at
similar sites on neighboring Montserrat in July 2000
(14% of phyllostomid bat captures).

In 2003, three males were captured-two in Bats
Cave (June 8) and one netted in fruit orchards in Chris¬
tian Valley (June 9). The Monophyllus taken in Bats
Cave were found in a small side tunnel that opened
in the floor of the right-hand chamber where no other
species were found. This small descending tunnel was
approximately 1 m wide and 1.5 m high, at least 30
m long, and may have had a separate opening to the

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Occasional Papers, Museum of Texas Tech University

outside as evidenced by a slight breeze in that tunnel.
At least 20 individual Monophyllus were observed
in this small space. Three captured males had testes
lengths of 5,4, and 3 and weighed 15.5, 13.6, and 15.4,
respectively.

Brachyphylla cavernarum cavernarum
Gray 1834

Specimens examined (114).-St. Paul Parish:
Bats Cave, 1.5 km E English Harbour Town, 23 m,
17°00 / 52 // N, 6r45'01"W, 20 (16 TTU, 4 UNSM);
Bats Cave, 2 mi. E Falmouth, 5 (2 FMNH, 3 UF); near
Bats Cave, English Harbor [= Harbour], 12 (NMNH);

1 mi. E English Harbor [=Harbour], 3 (1 KU, 2 MSU).
Parish Unknown: no specific locality, 74 (1 BMNH,
73 NMNH).

Specimens captured/released (105).-St. Mary
Parish: Boggy Peak Road, 1; Cades Bay Plantation, 6;
Christian Valley Agriculture Station, 1; Wallings Res¬
ervoir, 2. St. Paul Parish: Bats Cave, 95 (Fig. 1).

Miller (1913a; see also Koopman 1968) was
the first author to report Brachyphylla cavernarum
from Antigua. Swanepoel and Genoways (1978)
examined 20 specimens from the island as part of
their systematic review of the genus. Swanepoel and
Genoways (1978) assigned the material from Antigua
to the nominate subspecies, which has a type locality
of St, Vincent. Table 1 presents the length of forearm
and seven cranial measurements for 12 males and
9 females from Antigua. The only measurement in
which there was a significant difference between the
sexes was mastoid breadth, with males being larger
(P<0.05). In general, males were larger in all other
cranial measurements except postorbital constriction
in which the sexes were similar. Females had longer
forearms. The measurements of our sample from An¬
tigua match closely those samples of B. c. cavernarum
presented by Swanepoel and Genoways (1978).

Sex ratio data for Brachyphylla hand-captured
in 2000 in Bats Cave suggest near male-female par¬
ity (48 males: 44 females). This male-female balance
was mirrored by a 5:4 male-female capture ratio while
mist netting in several foraging habitats located many
kilometers away from the cave. On 3 August 2000,
we observed a mixed male/female maternity colony of

Brachyphylla cavernarum numbering well over 20,000
animals in Bats Cave (independent estimates by three
observers). In contrast to the Brachyphylla maternity
colony on the neighboring island of Montserrat, the
Brachyphylla colony on Antigua was not separated
into male/female caves, nor was there any obvious
evidence of male-female territoriality. Animals were
observed in clusters ranging in size from 10 to 200
individuals. The clusters of animals occupied numer¬
ous pockets in the domed roof of each chamber of
the cave. Neonates and juvenile animals appeared to
be randomly located throughout the cave, giving no
indication of a “creche.” Numerous mid-air collisions
between individual Brachyphylla were observed, sev¬
eral resulting in the grounding of both animals. Due
to the apparent difficulty in taking flight from the soft
substrate (guano), most grounded bats climbed up onto
an adjacent rock surface before taking off. Of note, the
exposed upper surfaces of most rocks in lower half of
the cave were highly burnished, no doubt in response
to animal contact.

Bats Cave was visited on 8 June 2003, when the
majority of the females in the colony were carrying
pups. The females were located in the deeper, hotter,
more humid portions of the cave. Many of the males
were congregated near the entrance and in the outer¬
most chambers where the temperature was considerably
lower and ambient light made the bats clearly visible,
Five of the six females obtained in 2003 were lactat-
ing and were collected along with their infants. The
sixth female was non-reproductive. The testes length
of six adult males collected at this time averaged 6.3
(4-9). The length of forearm and weight of three male
and one female infant were 56.6, 51.6,51.8, and 52.2
and 25.6, 21.8, 20.6, and 20.7, respectively. A male
captured on 25 June 1958, had testes that were 5.0
long. Of 10 females collected and preserved from Bats
Cave on 29 April 1959, nine were pregnant. Some of
the embryos had been removed for other studies, but
the crown-rump length of remaining four embryos was
34, 36, 37, and 39. All 21 females collected on 1 May
1958 were pregnant, with one of the embryos measur¬
ing 37 in crown-rump length. None of the nine females
taken on 2 and 7 August 1903, the 3 females taken on
7 November 1903, or the 5 females from 20 November
1903 were pregnant. These combined reproductive
data suggest that a monestrus reproductive pattern
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
and 2000 capture/release) for 12 adult males was 66.2
(64.6-69.0) and 49.5 (41.8-56.0), and the same for six
adult females was 66.2 (64.0-68.0) and 46.4 (41.1-
49.4), respectively. Five lactating females captured in
early June 2003 weighed an average of42.7 (38.6-46.3),
whereas a non-reproductive adult female weighed 36.5.
Six adult males taken at this time weighed an average
of 43.4 (39.0-48.5).

Morton and Lindsay recorded roost emergence
behavior for Brachvphylla from Bats Cave in February
1994. They observed that a small number of bats began
to exit the cave to forage around 1845 h, whereas the
majority of the colony waited until after dark (1915 h)
to exit the cave. The emergence continued sporadically
through 1945 h.

Radio-tracking data for two male Brachvphylla
indicate that these robust bats travel nightly up to 22
km away from Bats Cave to forage in the moist, higher
altitude valleys and farm lands on the southwestern
side of the island. In addition, one of the wing-banded
Brachyphylla was recaptured at 14.3 and 15.4 km from
the roost on consecutive evenings (minimum-commut¬
ing distance of 60 km over two nights).

Artibeus jamaicensis jamaicemis
Leach 1821

Specimens examined (28).-St. John Parish:
Santa Maria Hill, 2+ mi. NE St. Johns, 4 (NMNH).
St. Mary Parish: Christian Valley, 2.1 km S Jennings,
32 m, 17°03'33"N, 61°51'44"W, 6 (TTU); Wallings
Reservoir, Wallings, 166 m, 17°02'04"N, 61°49'30"W,
6 (TTU). St. Paul Parish: English Harbor [= Harbour],
10(1 MCZ, 9 NMNH). Parish Unknown: no specific
locality, 2(1 FMNH, 1 MCZ).

Specimens captured/released (80).-St. John
Parish: Green Castle Agricultural Station, 7. St. Mary
Parish: Boggy Peak Road, 9; Cades Bay Plantation, 6;
Christian Valley Agriculture Station, 10; Claremont
Plantation, 11; Wallings Reservoir, 12. St. Paul Parish:
All Saints, 4; Swetes Dam, 16. St. Philip Parish: Col¬
lins, 1; Montpellier, 4 (Fig. 1; Table 1).

J. A. Allen (1890, see also G. M. Allen 1911) first
reported this species from Antigua based on an adult

male captured on 30 December 1889, and an adult fe¬
male captured on 23 April 1890. Genoways et al. (2001)
reviewed Antillean populations of the Jamaican fruit
bat based upon morphometries and presence/absence of
M3/m3 and Phillips et al. (1989) and Pumo et al. (1996)
presented genetic data for these populations. These
studies support the use of the subspecific name A. j.
jamaicensis for Antiguan populations. Table 1 presents
length of forearm and seven cranial measurements for
four male and five female A. jamaicensis from Antigua.
The sexes differed significantly only in postorbital
constriction in which females were larger on average.
Females were larger on average than males in all other
measurements as well except length of the maxillary
toothrow in which the sexes had the same mean.

Eighteen of the 29 males (62%) captured and
released in July 2000 exhibited scrotal testes. Of the 35
females captured and released in July 2000, five were
pregnant and 11 were lactating or post-lactation (45%
of the females were reproductively active). Six adult
males collected between 7-9 June 2003, had testes that
averaged 9 (6-11) in length. Of six females taken during
this same time period, three evinced no reproductive
activity, two were lactating, and one carried an embryo
that was 20 mm in crown-rump length. Two females
taken at Santa Maria Hill on 14-15 August 1980 were
lactating.

For individuals that were captured and released,
the average length of forearm and body mass for 12
adult males were 59.5 (57.0-63.0) and 40.9 (36.3-50.6),
respectively, and the same for 26 adult non-pregnant
females were 60.4 (57.0-62.0) and 42.5 (34.7-50.4),
respectively. The weights of three non-pregnant females
taken in early June 2003 were 34.7, 36.9, and 36.9,
whereas two lactating females weighed 39.7 and 39.9.
Six adult males, also taken in early June 2003, weighed
an average of 38.9 (35.5-43.2).

Although W. R. Forrest recorded in his field
notes that Artibeus was present in “Bats Cave” in 1918,
Artibeus was not observed there in 1994, 1998, 2000,
or 2003. However, Artibeus was commonly observed
in abandoned buildings and was collected at all mist-
netting localities in 1994, 1998,2000, and 2003. These
animals were observed hanging pendant from foliage
and from the ceilings/rafters of unused buildings, either
individually or in very tight clusters. The individuals

Occasional Papers, Museum of Texas Tech University

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captured at Santa Maria Hill were taken from a lime¬
stone overhang. In direct contrast to the raucous nature
of BrachyphyUa roosts, Artibens roosts typically remain
quiet unless the animals are disturbed by attempts to
capture them.

Of the eight individuals examined, seven were
missing both upper third molars and possessed both
lower third molars. The eighth individual was missing
both third upper molars as well as the lower left third
molar. The lower right molar was minute. On Antigua,
heavy tooth wear was noted only on a single female
Artibeus in 2000 (1% of captures) that was clearly a
very old animal.

Natalus stramineus stramineus Gray 1834

Specimens examined (8).-St. Paul Parish: Bats
Cave, 1.5 km English Harbour Town, 23 m, 17°00'52"N,
6r45'01"W, 1 (UNSM); English Harbor [= Harbour],
2 (MCZ). St. Philip Parish: Montpelier Cave, 1.2 km
E, 0.2 km S, 78 m, 17°02'36"N, 61°425'2"W, 3 (TTU).
Parish Unknown: no specific locality, 2 (MCZ).

Specimens captured/released (9).-St. Mary Par¬
ish: Christian Valley, 2. St. Paul Parish: Bats Cave, 7.

Goodwin (1959), in his revision of members of
the subgenus Natalus, restricted the type locality of
Natalus stramineus to Antigua rather than Lagoa Santa,
Minas Gerais, Brazil, as was done by earlier authors.
With this decision, which later was reconfirmed by
Handley and Gardner (1990), the nominate subspe¬
cies N. s. stramineus becomes the appropriate name to
apply to the population on Antigua. Goodwin (1959)
gives measurements for two males from Antigua in the
British Museum (Natural History) as follows: length of
forearm, 38.0,38.8; greatest length of skull, 16.6, 16.6:
condylobasal length, 15.5, 15.7; zygomatic breadth,
8.7, 8.8; breadth of braincase, 8.1, 8.0; interorbital
breadth, 3.4,3.5; breadth across upper molars, 5.9, 5.9;
length of maxillary toothrow, 7.4, 7.5. Table 1 presents
the length of forearm and seven cranial measurements
for three males and three females from Antigua. The
range of measurements of these topotypic individuals
fall within the range of a larger sample from Dominica
(Genoways et al. 2001).

Seven specimens were hand-collected in Bats
Cave, while two others were hand-collected in a large
rock crevice between very large boulders in Christian
Valley in 1994 (1600 h) (Fig. 1). In 2000, numerous
Natalus (n ~= 90+) were observed in Bats Cave, but
no young were observed. As is typical of the species,
each bat hung fully pendant a short distance (15-20
cm) from neighboring bats across the ceiling of Bats
Cave. In 2000, Natalus was again observed to utilize
height restricted portions of the cave ceiling along large
fissures that diverge laterally from the main chamber.
The apparent preference for these smaller side passages
along the middle and at the back of the cave may be
due to a variety of reasons including: (1) the slightly
higher relative humidity in these microclimates (85%
humidity), and (2) a desire to avoid the much larger
BrachyphyUa. Indeed, we witnessed a mid-air collision
between Natalus and BrachyphyUa that fractured the
right humerus of the smaller bat (UNSM 27900). When
funnel-eared bats began emerging around 1840 h, they
were flying low to the ground, but did not appear to have
made use of the main exit hole. The facultative use of
secondary exits may avoid collisions with BrachyphyUa
that also began to exit around this time. No Natalus
were observed in Bats Cave on 8 June 2003.

During the 1998 and 2003 surveys, a limestone
cave 600 m east of the disused Montpellier sugar fac¬
tory east of St. Philips was visited. This tunnel ran for
nearly 120 m with a relatively constant diameter of 1.5
to 2.0 m. At least five Natalus were observed in this
cave in 1998 and 12 to 15 animals were observed in
2003. The northeastern half of Antigua is dominated by
xeric habitats like those surrounding the Montpellier
Cave and Bats Cave. These caves are very important
from a management perspective as they provide criti¬
cal humid microenvironments that Natalus must have
across this xeric part of the island.

Length of forearm and body mass for two fe¬
males captured/released in 1994 and 2000 were 39.0
and 40.0 and 5.2 and 6.5, respectively. A female and
two males taken on 9 June 2003, weighed 4.4, 4.8,
and 5.0, respectively. The female evinced no gross
reproductive activity and the males had testes lengths
of 1.0 and 2.0.

Pedersen et al.-Bats of Antigua, Northern Lesser Antilles

11

Molossus molossus molossus (Pallas 1766)

Specimens examined (20).-St. Mary Parish:
Christian Valley, 2.1 km S Jennings, 32 m, 17°03'33"N,
61°51'44"W, 8 (TTU); Wallings Reservoir, Wallings,
166 m, 17°02'04"N, 61°49'30"W, 9 (TTU). St. Paul
Parish: English Harbor [= Harbour], 1 (NMNH). St.
Philip Parish: Mill Reef, 1 (AMNH). Parish Unknown:
no specific locality, 1 (NMNH).

Specimens captnred/re/cased (8).-St. Mary Par¬
ish: Cades Bay Plantation, 3; Claremont Plantation, 1;
Wallings Reservoir, 1. St. Paul Parish: Swetes Dam,
2. St. Philip Parish: outbuilding adjacent to Newfield
Moravian Church, 1 (Fig. 1; Table 1).

Miller (1913b; see also Miller 1924: 89) included
the first specimens of this species reported from Anti¬
gua in the material that he described under the species
name Molossus debilis and Koopman (1968) listed
two specimens from Antigua obtained by Clayton E.
Ray in 1963. The name M. debilis and a number of
others have been applied to this bat in the Antillean
islands. It is now believed that the most appropriate
name to apply to these bats is Molossus molossus ,
which is a widespread Neotropical species. Husson
(1962) restricted the type locality of M molossus to
the island of Martinique, which lead Dolan (1989) to
apply the name M. m. molossus to this species in the
Lesser Antilles, Table 1 presents the length of forearm
and seven cranial measurements of five males and five
females from Antigua. Males were significantly larger
than females at the P < 0.01 level for greatest length of
skull and at the P < 0.05 level for condylobasal length
and mastoid breadth. Males were larger than females on
average for length of forearm and zygomatic breadth,
whereas the sexes had the same mean values for post¬
orbital constriction, length of maxillary toothrow, and
breadth across upper molars.

Between 29 July and 1 August 2001, three preg¬
nant females and one lactating female were captured
and released. Eight of 11 females netted 7-9 June 2003,
were pregnant carrying single embryos, with the fol¬
lowing crown-rump lengths: 6, 7, 7, 8, 9, 10, 10, and
16, Four males captured at this time had testes lengths
of 3, 4, 4, and 6.

During the period 28 July - 2 August 2000, the
length of forearm and body mass for a single adult
male captured/released were 38.0 and 14.0, whereas
the average length of forearm and body mass for three
adult females were 35.3 (34.0-36.0) and 10.0 (9.6-10.8);
three pregnant females, 36.0 and 13.8 (12.6-14.8); and
a single lactating female 37.0 and 11.9, respectively.
The three non-pregnant females obtained in early June
2003 had weights of 9.1, 10.1, and 10.2, whereas the
eight pregnant females weighed an average of 11.0
(10.0-12.2). Five males collected at this time weighed
an average of 11.4 (10.6-13.1).

Tadarida brasiliensis antillularum
(Miller 1902)

Specimens examined (28),-St. Mary Parish:
Wallings Reservoir, Wallings, 166 m, 17°02'04"N,
61°49'30"W, 16 (TTU). St. Paul Parish: English Harbor
[= Harbour], 12 (NMNH).

Specimens captured/released (17)—St. Mary Par¬
ish: Flat Top Hill, 9. St. Philip Parish: Newfield, 8.

Shamel (1931) reported 21 specimens of this
species from Antigua under the name Tadarida antil-
lularum. Miller (1902) had described this taxon with
a type locality at Roseau, Dominica. Schwartz (1955)
reduced this taxon to a subspecies in the widespread
Tadarida brasiliensis , with a geographic range extend¬
ing from Puerto Rico to St. Vincent (Genoways et
al. 2001). Table 1 presents the length of forearm and
seven cranial measurements for five males and five
females from Antigua. Female Tadarida were found
to have a significantly longer forearm than males
at the P < 0.01 level, whereas males had a signifi¬
cantly longer condylobasal length than females at the
P < 0.05 level. The genders did not differ significantly
in any of the other measurements, but males were
somewhat larger on average for greatest length of
skull, postorbital constriction, and mastoid breadth, and
females were larger on average for zygomatic breadth.
The sexes averaged the same for length of maxillary
toothrow and breadth across upper molars.

During 1994, Morton and Lindsay examined
seventeen specimens that had been hand-collected
during daylight hours. Eight specimens of Tadarida

12

Occasional Papers, Museum of Texas Tech University

were eollected from under the roofing of a derelict
outbuilding at the Moravian Church in the town of
Newfield (6 males, 2 females; 1830 h) on 2 February.
Nine other animals were extracted from tiny crevices in
a rock overhang near Flat Top Hill (3 males, 6 females;
1915 h) (Fig. l)on 16 January. On the night of 6 June
2003, five male and four female Tadarida were netted
over water in the Wallings Reservoir and on 9 June two
males and five females were taken in the same place.
In 2003, the water behind the dam was greatly reduced,
but the pool was still at least 20 m wide and 50 m long.
The specimens collected in 1918 by W. K. Fisher from
English Harbour were all females.

Of the nine females netted in early June 2003,
seven evinced no gross reproductive activity, one was

lactating, and one carried a single embryo measuring
5.0 in crown-rump length. Seven males taken during
this time had testes that averaged 3.9 (3-5) in length.

Average length of forearm (mm) and body mass
for nine males captured and released in 1994 were 38.6
(37.0-40.0) and 9.4 (8.7-10.5), respectively, whereas
eight females were 38.7 (38.0-40.0) and 9.2 (7.9-
11.1). Seven nonpregnant females captured in June
2003 weighed an average of 9.6 (8.8-10.4), a lactating
female weighed 9.1, and a pregnant female weighed
9.5. Seven males taken at this time weighed an average
of 9.0 (7.9-9.8).

Discussion

In theory, the number of species found on islands
along an archipelago is correlated with the size (area)
of each island, but this effect is ameliorated by distance
from a source area such as a continent. The number
of species occurring on an island also is dependent
on the diversity of habitats available, which in most
cases is directly affected by elevation of the island.
Increased elevation usually results in increased rain¬
fall and a more diverse vegetation (MacArthur 1972).
For example, Morgan and Woods (1986) studying the
whole West Indian mammalian fauna found that 69%
of the variance in species diversity could be explained
by area of the islands alone. They concluded that “The
remaining 31 % of the variance must be dependent upon
other variables such as habitat diversity and distance
from source areas.”

Following models that have been applied to
amphibians and reptiles (Preston 1962a, 1962b), birds
(Hamilton et al. 1964), and West Indian bats or mam¬
malian fauna (Morgan and Woods 1986; Griffiths and
KJingener 1988), we constructed a species-area curve
for the bat fauna throughout the Antilles (Fig. 2). Under
this general model, several islands including Antigua
fall below the regression-line relative to other islands
of their size. These deviations amount to the “absence”
of species from their respective chiropteran faunas.
These particular islands may simply be under-sampled
(Saba) or perhaps these islands may not provide suf¬

ficient habitats to support bat diversity, despite their
size (Antigua, St. Eustatius).

Antigua is situated at the northern end of what
Genoways et al. (1998, 2001) termed the Lesser An¬
tillean Faunal Core. However, the paucity of chirop¬
teran taxa on Antigua (notably the absence of Arc/ops
niehollsi, Chiroderma improvisum, Stumira thomasi,
Eptesicus guadeloupensis , and Myotis dominicensis)
distinguishes Antigua from neighboring Guadeloupe
(Genoways and Baker 1975; Genoways and Jones
1975; Jones and Genoways 1975; Jones and Phillips
1976). Neither species of insectivorous bat ( Eptesicus
guadeloupensis and Myotis dominicensis) has been
reported from Antigua, or from neighboring Montserrat
despite extensive efforts that span 20 years on those
islands (Adams and Pedersen 1999; Jones and Baker
1979; Morton and Lindsay 1994; Morton and Fawcett
1996; Pedersen et al. 1996; Pierson and Warner 1990).
However, it should be remembered that for most is¬
lands, Myotis is known only from a single specimen
or from a small series. Following an extensive earlier
survey (Baker et al. 1978), the genus Myotis was first
reported from Guadeloupe based on only two speci¬
mens in 1992 (Masson and Breuil 1992). The other
insectivorous species missing from both Antigua and
Montserrat is Eptesicus guadeloupensis , which is cur¬
rently known only from Guadeloupe.

Pedersen et al.-Bats of Antigua, Northern Lesser Antilles

13

Figure 2. Species/area curve for the chiropteran fauna in the Antilles. Data are from Genoways et al. (2001) with minor modi¬
fication. Linear regression of log-transformed data: y = 0.17x + .49 (R-squared = 0.81). For argument (see text), Antigua has
been represented twice with the extant compliment of seven species of bats (closed circle) and with it’s compliment increased
by three additional species (Arrow: Square).

Conversely, three species of fruit bat -Ardops
nichollsi , Stnrnira thomasi (Genoways 1998; Pedersen
et al. 1996), and Chiroderma improvisum (Jones and
Baker 1979; Pierson and Warner 1990)-have been
reported from both of Antigua’s neighbors, Montserrat
(50 km WSW) and Guadeloupe (75 km SSE; Baker et
al. 1978; de la Torre and Schwartz 1966), but have not
been found on Antigua. Most notable by its absence
is Ardops nichollsi , a common tree-roosting fruit bat
found throughout the Lesser Antilles that is quite com¬
mon on the smaller nearby islands of Montserrat (ap¬
proximately 15% of frugivore captures; Pedersen et al.
1996) and Nevis (Pedersen et al. 2003). Despite efforts
to sample nearly identical habitats on Montserrat and
Antigua in 2000, no Ardops have been observed nor
collected on Antigua.

Part of the explanation for the differences in the
chiropteran fauna of Antigua as compared to those of
Guadeloupe and Montserrat may lie in the elevations
of the islands-Guadeloupe, area 1779 km 2 , maximum
elevation 1465 m; Antigua, 279 km 2 , 405 m; Montser¬
rat, 102 km 2 ,910m. The much lower average elevation
of Antigua would have resulted in less diversity of the
vegetation of the island. As Loveless (1960) stated,
the island originally may have been forested, but this
would have been a low scrub forest with limited areas
of Evergreen Seasonal forest. This forest type probably
would not provide a source of fruit for the fruit-eating
bats throughout the year, thus their absence may be
attributed to the lack of diversity in the flora of the
island. The apparent absence of several insectivorous
species of bats also may relate to this same issue. Again
with the reduced diversity of flora on the island there

14

Occasional Papers, Museum of Texas Tech University

should be a resulting reduction in the diversity of the
insect fauna, which consequently may not support a
diversity of insectivorous bats,

The missing species of bats on Antigua also may
lie in the fossil record of the island. Steadman et al.
(1984) and Pregill et al. (1988) reported on three species
of bats in the fossil record of Antigua that no longer
occur on the island Pteronotus parnellii , Mormoops
blaimUlii , and Phyllonycteris major -from deposits
that range from 4300 years B.P. at the bottom to 2560
years B.P. at the top. They also reported four species in
the fossil record that still live on Antigua Brachyphylla
cavenmnim, Natalns stramineus , Tadarida brasiliensis,
and Molossus mo loss us. Finally, they described a man¬
dible from a glossophagine bat, possibly Glossophaga
ox MonophyHits, the latter being more likely as it is still
extant on Antigua.

Only recently has Pteronotus parnellii been
discovered in a living chiropteran fauna on a Lesser
Antillean island-St. Vincent (Vaughan and Hill 1996).
Historically, P. parnellii could have reached Antigua ei¬
ther from the south or from the northwest, but Vaughan
and Hill concluded that the population on St. Vincent
was more closely related to South American mainland
populations than to those that occur in the Greater
Antilles (Smith 1972), a conclusion that has been
subsequently supported by Lewis-Oritt et al. (2001)
who used molecular data to show that all populations
ofP parnellii in the Antilles would have originated in
northeastern South America.

Mormoops blamviUii has occupied a broader
geographic range in the past than at the present time,
with fossil records from Abaco, Exuma, and New
Providence in the Bahamas and Anguilla and Barbuda
in the Lesser Antilles (Koopman 1951; Morgan 2001).
Antigua is far removed from the present geographic
ranges of Mormoops blainvillii and the fossil species
Phyllonyeteris major whose eastern-most locality is
Puerto Rico.

The chiropteran fauna of Antigua raises some
interesting questions for island biogeographers. What
is the appropriate slice of time that should be used
w'hen constructing species-area curves? Using only the
data from the last 200 years, the chiropteran fauna of
Antigua appears unbalanced, but if data from the last
4000-5000 years are used, the chiropteran fauna seems
to fit the predicted species-area curve. How' should hu¬
man impact be factored into species-area curve analy¬
ses? Steadman et al. (1984) attributed the loss of the
three species of bats and other vertebrates on Antigua
to “human-caused environmental degradation in the
past 3500 yr.” However, given the accelerated rate of
development and deforestation on these small islands
in the last 25 years, how should conservation officers
best utilize species-area curves in their management
decisions? How can elevation be factored into spe¬
cies-area curves? Should insectivorous and frugivorous
guilds be treated separately? Clearly, there needs to be
a re-evaluation of how to factor-in the entire profile of
an island into a biogeographical analysis.

Pedersen et al.-Bats of Antigua, Northern Lesser Antilles

15

Acknowledgments

We wish to thank the following for financial and
materiel support of the project: Totten Trust, Mariah
Fund, Hagan Fund of the University of Nebraska
State Museum, Nature Conservancy, Environmental
Awareness Group Biodiversity Programme-Antigua,
Island Resources Foundation-Antigua, British Airways,
Jersey Wildlife Preservation Trust, Flora & Fauna
International, and the British High Commission-An-
tigua. We thank local authorities for their assistance
in arranging collecting permits and access to public
lands: M. Henry, Chief Forestry Officer, Ministry
of Agriculture, Antigua; G. Cooper of the Environ¬
mental Awareness Group, Antigua; K. Lindsay of the
Island Resources Foundation, Antigua. The patience
and courtesy extended to us by the Estate owners on
Antigua are gratefully acknowledged. We also wish
to thank the following for their assistance in the field:
D. Black, J. Burton, O, Davis, S. Daane, N. Gricks, H.

Pitcher, T. Richards, and B, Smith. Curatorial support
was provided by the Division of Zoology, University
of Nebraska State Museum, with special thanks to T.
Labedz. A. Fox, Staff Artist for the University of Ne¬
braska State Museum, prepared Figure 1. We also wish
to thank the following curators and collection managers
for allowing us access to specimens housed in their
research collection: N. B. Simmons, American Museum
of Natural History; P Jenkins, British Museum (Natu¬
ral History); B. D. Patterson and L. R. Heaney, Field
Museum of Natural History; R. M. Timm, Natural His¬
tory Museum, University of Kansas; J. M. Chupasko,
Museum of Comparative Zoology, Harvard University;
L. Abraczinskas, The Museum of Michigan State
University; R. W. Thorington, Jr., National Museum
ofNatural History; R. J. Baker, Texas Tech University;
C. McCaffery, Florida Museum of Natural History,
University of Florida.

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18

Occasional Papers, Museum of Texas Tech University

Addresses of authors:

Scott C. Pedersen

Department of Biology and Microbiology’

South Dakota State University
Brookings, SD 57007
Scott_Pedersen@sdstate. edu

Hugh H. Genovvays

W436 Nebraska Hall

University of Nebraska State Museum

Lincoln, NE 68588

hgeno ways 1 @unl. edu

Matthew Morton

c/o 22 Kens a I Avenue
Bedminster

Bristol, BS3 4QY United Kingdom
matthewnmorton@hotmail.com

Vicki J. Swier

Department of Biological Sciences and Museum
Texas Tech University
Lubbock, TX 79409
vicki_swier@hotmail.com

Peter A. Larsen

Department of Biological Sciences and Museum
Texas Tech University
Lubbock, TX 79409
plarsenOl @yahoo. com

Kevel Lindsay

Island Resources Foundation
1718 P Street, N.W., 4T4
Washington, DC 20036
klindsay@irf.org

Rick Adams

Department of Biological Sciences
University of Northern Colorado Ross Hall
501 20th Street Greeley, CO 80639
rick. adams@unco. edu

Jon Appino

Department of Biology and Microbiology
South Dakota State University
Brookings, SD 57007

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