Presence of Stenoderma rufum beyond the Puerto Rican bank

2007·0.6 MB·English

by  Kwiecinski, Gary George, 1952-

TEXAS TECH UNIVERSITY
Natural Science Research Laboratory
Occasional Papers
Museum of Texas Tech University Number 266 1 June 2007
Presence or Stenoderma rufum Beyond the Puerto Rican Bank
GaryG. Kwiecinski and William C. Coles
Abstract
Until now the red fig-eating bat, Stenoderma rufum, had been known only from the
Caribbean islands of Puerto Rico, St. Thomas, and St. John. The capture of one specimen on
St. Croix is noteworthy because this is the first reported capture of this species beyond the
Puerto Rican bank. The female captured was post-lactating and pregnant, suggesting this is
not an isolated individual, but a breeding population on St. Croix is more likely. St. Croix’s
distinct origin and independence from the northern Virgin Islands indicate S. rufum most likely
arrived by over water dispersal to St. Croix from adjacent islands (St. Croix is 68 km south of
St. John and St. Thomas and 101 km southeast of Puerto Rico). When morphological characters
and habitat characteristics are compared between the St. Croix Stenoderma with female S. r.
rufum from St. Thomas and St. John and S. r. darioi from Puerto Rico, none clearly align the
St. Croix specimen with either subspecies, except pelage color is consistent with S. r. rufum.
Key words: cranial characters, external morphology, St. Croix USVI, Stenoderma rufum
Introduction
The red fig-eating bat, Stenoderma rufum, is the type locality indicated as “unknown” due to its
known from the Caribbean island of Puerto Rico and doubtful origin. When Peters (1869) visited the Paris
the northern United States Virgin Islands (USVI) of St. Museum and examined the type represented by a dam¬
Thomas and St. John (Fig. 1) but not from the south¬ aged skin and no skull, he was deceived by the faulty
ern USVI of St. Croix (Hall and Bee 1960; Hall and figure of Geoffroy and named the bat a Vampyrops.
Tamsitt 1968; Genoways and Baker 1972; Koopman Peters (1876) subsequently examined the type of Ar-
1975; Timm and Genoways 2003). Stenoderma was tibeus undatum Gervais (skull only, skin thought to be
first described by Etienne Geoffroy St.-Hilaire (1818) lost) in the anatomical collection at the Paris Museum.
as “le Stenoderme roux” based on a single specimen, The similarity of the skull of A. undatum and the
the type locality not known, but it originally was be¬ skull figured by Geoffroy led Peters to conclude the
lieved to have come from Egypt. Since the one known skull of A. undatum was actually the missing skull of
specimen resembled New World bats rather than bats Stenoderma rufa. This established the identity of the
known from Egypt or the Old World, Demarest (1820) species, but not its habitat. The supposition that the
used Stenoderma rufa in his original description and genus and species would be rediscovered in the West
2 Occasional Papers, Museum of Texas Tech University
Figure 1. Current geographic distribution of Stenoderma rufum in the West Indies includes S. r. darioi from Puerto Rico
and S. r. rufum from the United States Virgin Islands of St. Thomas, St. John and St. Croix. The inset map and its arrow ap¬
proximate locations of Puerto Rico and Virgin Islands in the West Indies. The United States Virgin Islands (slanted lines) are
defined by the boundary lines to their west to distinguish their territory from Puerto Rico (grey) and to the east to distinguish
them from the British Virgin Islands (black).
Indies was accepted by scientists studying American and Bee 1960). Differences in cranial and external
Chiroptera (Allen 1911). The distribution and status measurements led Hall and Tamsitt (1968) to classify
of this taxon remained quiescent until Anthony (1918, specimens from Puerto Rico as a new subspecies, S.
1925) discovered sub-Recent fossil remains in caves rufum darioi, and Choate and Birney (1968) charac¬
of Puerto Rico. No additional living specimens were terized fossil remains from Puerto Rico as a separate
collected until 1943, when Harry A. Beatty obtained subspecies, S. r. anthonyi. Since confirmation of the
a specimen on St. Thomas, US VI, but at the time the existence of this species (Hall and Bee 1960), numer¬
specimen was not identified to species. The St. Thom¬ ous authors have captured these bats on Puerto Rico
as specimen was housed at the U. S. National Museum and various aspects of its natural history and biology
of Natural History (NMNH), Washington, DC, among are slowly being revealed (e.g., Genoways and Baker
unidentified specimens until Dr. Charles O. Handley, 1972; Scogin 1982; Willig and Bauman 1984; Willig
Jr. recognized it as S. rufum (Hall and Tamsitt 1968). and Gannon 1991; Gannon and Willig 1992). We
Three living individuals captured in 1957 on St. John, describe a post-lactating, pregnant female specimen
USVI (Hall and Bee 1960), confirmed the contempo¬ of S. rufum captured on St. Croix, USVI (Fig. 2) and
rary existence of this species. These three specimens compare its external and skull measurements to those
were deemed indistinguishable from the holotype and from female S. rufum from Puerto Rico.
they established St. John as the type locality (Hall
Materials and Methods
On the evening of 10 June 2004, we set three proximately 110-140 cm of rain a year. Creque Ghut
mist nets (38 mesh; Avinet, Dryden, NY) near the is one of a very few locations on St. Croix with per¬
Creque Reservoir, in the West End Quarter, Frederik- manent running water (in the “swimming hole” area),
sted District, St. Croix, USVI. Creque Ghut, paral¬ although most of its length is intermittent stream. The
leled by Creque Dam Road, winds up a long, steeply over-story is dominated by kapok (Ceibapentandra),
sloped ghut through secondary growth rain forest sandbox (Hura crepitans), and mango (Mangifera in-
(Beard 1949). This semi-evergreen rainforest covers diccf). The under-story is dominated by tan-tan (wild
the northwestern third of St. Croix and receives ap¬ tamarind, Leucaena leucocephala). One mist net was
Kwiecinski and Coles-Stenoderma rufum from St. Croix 3
Figure 2. Photograph of live female Stenoderma rufum captured on St. Croix, 10 June 2004.
set along the length of the top of reservoir dam. A were fixed in formalin for a minimum of 48 hrs at
second mist net was set perpendicular to the dam, 4-10°C before being transferred to 70% ethanol and
parallel to and between the reservoir and Creque transported back to The University of Scranton.
Dam Road. The third net was set approximately 50 These became voucher specimens deposited at
m east (upstream) of the reservoir, in the ghut lead¬ the Museum of Texas Tech University, Lubbock,
ing to the reservoir, and at a location known as the Texas. The captured S. rufum was assigned catalog
“swimming hole.” The locality is identified as: St. number TTU-M104503, and the skull was cleaned
Croix; Frederiksted District; Northside A; Creque (dermestid beetles) and subsequently measured.
Reservoir; 0.25 km south, 0.1 km east Frenchman Measurements were taken following Hall (1946),
Hill; 17°44'45"N, 64°52'32"W. Elevation of the except the greatest length of the skull included the
netting site ranged from about 65 m (dam) to 100 incisors. The single specimen of S. rufum captured
m (swimming hole). on St. Thomas in 1943 by Harry A. Beatty was not
previously measured. This specimen is a parous
Mist nets were open from approximately female, non-reproductive at capture, stored in al¬
1845 to 2230 hr. Netted bats were held until all cohol at the National Museum of Natural History
nets were dismantled to reduce their stress and to (NMNH) as 271210, and no data or measurements
prevent recapture. After all nets were closed and were reported. At our request, the skull was ex¬
secured, the bats were weighed (g), the forearm tracted and prepared and we report cranial and
measured (mm) as the distance from the olecranon external measurements. Our measurements of the
process to the tip of the carpals with the wing in St. Croix and St. Thomas S. rufum were compared
a retracted position, and each bat was examined to other measurements of female S. rufum from
for ectoparasites, age, reproductive condition, and Puerto Rico and St. John that were currently avail¬
health status before being either released or held for able from literature.
further processing. Bats held for further processing
4
Occasional Papers, Museum of Texas Tech University
Results
The weather at the netting site on St. Croix, 10 mm (mean + SD): greatest length of mandible, 14.64
June 2004, was partly cloudy with occasional showers, ±0.15; height of coronoid process 7.96 ± 0.26 (Choate
temperature was 27°C and humidity was 93% (Kes¬ and Birney 1968). These fossil mandibular lengths
trel 3000 Pocket Weather Meter, Nielsen Kellerman, were greater than those of contemporary Puerto Rican,
Chester, PA) at 1830 hr prior to a rain shower. After St. Croix, St. John, and St. Thomas specimens, where¬
a 5 min down-pour, while opening nets at 1830-1835 as the height of the coronoid process was larger from
hr, humidity increased to 99-100% and remained fossils than from St. Croix, St. John, and St. Thomas
elevated for the remainder of our net sets. At 2230 specimens but comparable to S. r. darioi (7.90 ± 0.04)
hr, temperature was 25.5°C. From ca. 1830 to 2230 from contemporary Puerto Rican specimens.
hr we captured 13 Artibeus jamaicensis, 2 Molossus
molossus, and 1 Stenoderma rufum. No bats were The total length of the body and the length of the
captured after 2130 hr. The S. rufum was captured in forearm (67.0, 50.21 mm, respectively) of the female
the second net parallel to and between the reservoir St. Croix Stenoderma are similar to the average value
and the road, elevation 68 m. This bat was flying from found for Puerto Rican females (67.8 mm, 49.7 mm)
the road to the reservoir. and larger than values for the female St. John specimen
(65.0 mm; 47.0 mm). No total body length was avail¬
The captured Stenoderma rufum was post- able for the St. Thomas specimen but forearm length
lactating (fur worn around obvious mammary gland (50.18 mm) was similar to the St. Croix specimen.
nipples) and pregnant. The coloration of this St. Croix The length of the foot of the St. Croix specimen (13
Stenoderma was tan to light brown, with a slight rust mm) is considerably smaller than the length of the foot
color to hairs of dorsum of forelimbs, hindlimbs, and from the St. John (16 mm) and St. Thomas (15 mm)
uropatagium, whereas the undersides appeared to specimens, but not different from the average value
contain more grayish than tan or brownish color and for female Puerto Rican specimens (13.8 ±0.37 mm).
no hint of reddish coloration. In a hand-written note There was no difference in the length of the ear from
submitted with the St. Thomas specimen to NMNH, St. Croix, St. John, and St. Thomas specimens (18 mm)
Harry A. Beatty described color “in the unsoaked and the average value from Puerto Rican specimens
condition is brown.” The alcohol soaked specimen (17.8 ±0.20 mm).
presently appears light brown to tan, but the color
may have been affected by leaching of pigment while For eight characters (total length, length of ear,
stored. For the St. Croix specimen, there were two, length of forearm, greatest length of skull, condylo-
bilaterally symmetric, white spots, one located where basal length, mastoid breadth, zygomatic breadth,
the wing joins the body and another longer, crescent¬ and maxillary tooth row), measurements (Table 1)
shaped white spot directed anteriorly and ventrally to from the St. Croix specimen were greater than values
each ear. There was also a ring of white fur surround¬ from the St. John specimen, but were comparable to
ing the mouth (Fig. 2). These white areas were best St. Thomas and Puerto Rican specimen values. Five
viewed from lateral and ventral views. characters (postorbital constriction, breadth across
upper molar 2, greatest length of mandible, depth of
The greatest length of mandible and height of skull, and height of coronoid process) were similar in
coronoid process (in mm), respectively, of the St. values between St. Croix, St. John, St. Thomas, and
Croix (13.49, 7.74) and St. Thomas (14.25, 7.47) Puerto Rican specimens. The breadth of the braincase
specimens are within ranges for the species (Table 1). was similar in St. Croix and St. Thomas specimens, but
These values are comparable to values for St. John and greater than the St. John and Puerto Rican specimens.
contemporary Puerto Rican specimens, but less than The basal length of the skull was similar in St. Croix
fossil Puerto Rican specimens. Fossil lower jaws of and St. Thomas specimens but greater than the St. John
five female (presumed) S. r. anthonyi from sub-Recent specimen (no comparable measurements available for
deposit in Cueva de Clara, Puerto Rico, measured in Puerto Rican specimens).
Kwiecinski and Coues-Stenoderma rufum from St. Croix
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6 Occasional Papers, Museum of Texas Tech University
Discussion
The finding of Stenoderma rufum on St. Croix sic-Cretaceous boundary and separated by rising sea
has increased the known geographic range for this spe¬ levels and tectonic contractions (Larue 1994), whereas
cies and it may be more extensive than that currently St. Croix has been emergent for the last 3-4 ma only
known. The genera Stenoderma, Ardops, Phyllops, (Speed 1989; McLaughlin et al. 1995). St. Croix’s
and Ariteus, all of which have a white shoulder spot distinct geological origin and independence from the
and shortened rostrum, are parapatric genera that northern Virgin Islands indicates that bat populations
form the Antillean endemic subtribe Stenodermatina most likely arrived by over water dispersals to St.
believed to have evolved from a single, ancestral in¬ Croix from adjacent islands, similar to that espoused
vader, that subsequently radiated and diversified on by Hedges (1996), and not by vicariance (Rosen
West Indian islands before populating the mainland 1975) or by a combination of dispersal and vicariance
(Baker and Genoways 1978; Baker et al. 2003; Ge- (Iturralde-Vinent and MacPhee 1999).
noways et al. 2005). Because the S. rufum we found
was pregnant and post-lactating, a reproductive popu¬ The location of the type locality is unknown, but
lation has been found off the Puerto Rican bank for the name Stenoderma rufum was restricted to speci¬
the first time. Finding a pregnant female in June was mens from St. John because they are indistinguishable
not surprising because this species is asynchronously from the holotype in pallor of pelage (Hall and Bee
polyestrous and pregnancy could occur any time of 1960). The dorsal coloration of S. r. rufum from St.
year (Gannon and Willig 1992). John are described as tan to buckthorn brown with
a reddish hue and underparts that are paler and less
The significance of this record is that it is the reddish and indistinguishable from the holotype (Hall
first for the genus Stenoderma beyond the Puerto Ri¬ and Bee 1960). Color of the dorsum of Puerto Rican
can bank. The Virgin Islands form two quite distinct S. r. darioi varies from tan to dark chocolate brown or
geographical units. Vieques, Culebra, St. Thomas, St. Dresden brown, whereas color of the ventrum is paler
John, Tortola, Virgin Gorda, Anegada, and associated (Jones et al. 1971; Genoways and Baker 1972). The
smaller cays, collectively are known as the Northern dorsum of the St. Thomas specimen was described
Virgin Islands, whereas St. Croix and its smaller cays as “brown” and its current alcohol soaked condition
form the Southern Virgin Islands (Fig. 1). The North¬ appears light brown to tan. The dorsum of the St.
ern Virgin Islands are on the Puerto Rican bank, which Croix specimen was light brown to tan, not chocolate
formed a land extension of Puerto Rico until about brown, lighter than buckthorn brown, and appeared as
10,000 years ago and is known as the Virgin Island a variant similar to St. John specimens; therefore, all
Platform (Heatwole and Mackenzie 1967). Southeast Virgin Island specimens are similar in coloration and
of the Virgin Island Platform, St. Croix is a pinnacle of lighter in color than Puerto Rican specimens.
an east-trending submarine ridge, the St. Croix Plat¬
form. St. Croix and its associated cays, approximately The habitats of the two subspecies of Steno¬
68 km south of St. John and St. Thomas and 101 km derma rufum differ in that S. r. rufum previously has
southeast of Puerto Rico, are flanked by the Caribbean been found in dry arborescent vegetation on St. John
Sea Basin (specifically, the 4,500 m deep Virgin Island and S. r. darioi typically has been found in dense rain
Basin) and to the southeast by Aves Ridge. The tec¬ forest in Puerto Rico (Hall and Tamsitt 1968). The
tonic complex of St. Croix has no evident equivalent St. Croix S. rufum was found in secondary rain for¬
on islands of the northeastern Caribbean (Speed 1989). est, whereas the St. Thomas specimen was captured
Volcanigenic sediments of comparable age on Puerto along a “streamway” (handwritten notes of Harry A.
Rico and the northern Virgin Islands have not had Beatty) in habitat that is dry scrub and arborescent
similar depositional, deformational, or metamorphic vegetation.
histories as those of St. Croix (Speed 1989; Speed
and Joyce 1989; Speed and Larue 1991; Larue 1994). The cranium of the St. Croix specimen is larger
Puerto Rico and the northern Virgin Islands emerged than the crania of females from St. John but compa¬
from submarine volcanic mountains near the Juras¬ rable to the cranium of the female specimen from St.
Kwiecinski and Coles-Stenoderma rufum from St. Croix 7
Thomas and female specimens from Puerto Rico. St. Croix Stenoderma to S. r. rufum. If secondary
Based on available measures for female S. rufum, sexual dimorphism holds for the St. Croix Steno¬
the overall size of external characters and cranial derma, then males would be expected to be smaller
characters of the St. Croix and St. Thomas specimens than females, as this was the finding for Puerto Rico
are more similar to female Puerto Rican S. r. darioi (Gannon et al 1992) and for St. John (Jones et al.
than to St. John S. r. rufum. For example, the crania 1971) specimens.
of Puerto Rican specimens are longer than those
from St. John and the length of the crania of the St. It had been predicted that St. Croix would lack
Croix and St. Thomas specimens are longer than the some of the faunal elements present on the other
cranium of the St. John female but comparable in Virgin Islands and absent in the Lesser Antilles and
length to those of females from Puerto Rico. Size of for bats, this was true only for Stenoderma rufum
entire individual, external characters, size of cranium, (Koopman 1975). St. Croix can now claim to have
and shape of skull do not differ radically between the same bat fauna as the other Virgin Islands, except
the two subspecies, but pelage color is consistently for Tadarida brasiliensis. The only records of the
different (Hall and Tamsitt 1968). The St. Croix and widespread T. brasiliensis from the Virgin Islands
St. Thomas specimens are both similar in pelage and are netted specimens from St. John (Hall and Bee
are consistent with the Virgin Island subspecies S. r. 1960), and there are no data on roosting sites in the
rufum. Neither morphological characters nor habitat Virgin Islands for this species. S. rufum is the only
characteristics clearly align the St. Croix specimen bat species on the Virgin Islands not found in the
with either subspecies, but pelage is consistent with Lesser Antilles.
S. r. rufum, and therefore, we tentatively assign the
We are grateful to Patrick J. Kelly for his techni¬ on earlier drafts of this manuscript. We are grateful
cal help setting and monitoring nets while on St. Croix to Linda Gordon at NMNH for allowing us access
in June 2004. We are very appreciative of the expert to measure the skull and examine notes for the St.
curatorial assistance of Robert J. Baker and Heath Thomas specimen. This work was supported in part
Garner, Museum of Texas Tech University. Hugh by a Faculty Development Research Grant to GGK
H. Genoways provided expert advice and criticism from The University of Scranton.
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Addresses of authors:
Gary G. Kwiecinski William C. Coles
Biology Department Division of Fish and Wildlife
University of Scranton United States Virgin Islands Department of Plan¬
Scranton, PA 18510-4625 ning and Natural Resources
ggk3 01@scranton. edu Fredricksted, St. Croix, USVI00840
wcoles@vitelcom. net