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Mammalogy Papers: University of Nebraska State Museum Museum, University of Nebraska State
4-24-2007
BATS OF SAINT MARTIN, FRENCH WEST INDIES/SINT MAARTEN, NETHERLANDS ANTILLES
Hugh H. Genoways University of Nebraska-Lincoln, [email protected]
Scott C. Pedersen South Dakota State University, Brookings, SD, [email protected]
Peter A. Larsen Texas Tech University, [email protected]
Gary G. Kwiecinski University of Scranton, Scranton, PA
Jeffrey J. Huebschman University of Wisconsin-Platteville, Platteville, WI
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Genoways, Hugh H.; Pedersen, Scott C.; Larsen, Peter A.; Kwiecinski, Gary G.; and Huebschman, Jeffrey J., "BATS OF SAINT MARTIN, FRENCH WEST INDIES/SINT MAARTEN, NETHERLANDS ANTILLES" (2007). Mammalogy Papers: University of Nebraska State Museum. 76. https://digitalcommons.unl.edu/museummammalogy/76
This Article is brought to you for free and open access by the Museum, University of Nebraska State at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Mammalogy Papers: University of Nebraska State Museum by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Mastozoología Neotropical, 14(2):169-188, Mendoza, 2007 ISSN 0327-9383 ©SAREM, 2007 Versión on-line ISSN 1666-0536 www.sarem.org.ar BATS OF SAINT MARTIN, FRENCH WEST INDIES/SINT MAARTEN, NETHERLANDS ANTILLES
Hugh H. Genoways,1 Scott C. Pedersen,2 Peter A. Larsen,3 Gary G. Kwiecinski,4 and Jeffrey J. Huebschman5
1 University of Nebraska State Museum, W436 Nebraska Hall, University of Nebraska-Lincoln, Lincoln, NE 68588
ABSTRACT: Six species of bats have been previously reported from the Antillean island of Saint Martin/Sint Maarten—Artibeus jamaicensis, Brachyphylla cavernarum, Molossus molossus, Tadarida brasiliensis, Noctilio leporinus, and Myotis nigricans nesopolus. Our field research reported herein documents an additional three species of bats from the island for the first time—Monophyllus plethodon, Ardops nichollsi, and Natalus stramineus. Re-examination of the single voucher of Myotis nigricans nesopolus has led us to exclude this species from the fauna of Saint Martin/Sint Maarten. Based on our field research and the study of specimens housed in museum collections, we present information on the eight species of bats that we have documented as occurring on Saint Martin/Sint Maarten. The average rate of fruit bat captures on Saint Martin/Sint Maarten (0.92 bats per net-night— BNN) falls towards the lower end of the range (0.65-2.47 BNN) reported from nearby islands in the northern Lesser Antilles and below the range (2.20-5.93 BNN) reported for mainland populations of Neotropical fruit bats. We discuss possible causes of these de- creased population levels and we express some concerns about the future conservation status of the chiropteran fauna of the island.
RESUMEN: Murciélagos de Saint Martin, Indias Occidentales francesas/Sint Maarten, Antillas holandesas. Seis especies de murciélagos han sido registradas previamente para la Isla de Saint Martin/Sint Maarten, en las Antillas: Artibeus jamaicensis, Brachyphylla cavernarum, Molossus molossus, Tadarida brasiliensis, Noctilio leporinus y Myotis nigricans nesopolus. Nuestras investigaciones de campo ha permitido documentar por primera vez la presencia de tres especies adicionales para la Isla: Monophyllus plethodon, Ardops nichollsi y Natalus stramineus. La reexaminación de un único ejemplar de Myotis nigricans nesopolus nos llevó a excluir a esta especie de la fauna de Saint Martin/Sint Maarten. Nuestros estudios de campo y la examinación de ejemplares depositados en colecciones de museos nos ha permitido presentar información sobre las ocho especies de murciéla- gos que hemos podido documentar en Saint Martin/Sint Maarten. El promedio de captura de murciélagos frugívoros (0.92 murciélagos por noche - BNN) cayó hacia el extremo mas bajo del rango (0.65-2.47 BNN) reportado para islas cercanas al norte en las Antillas Menores, y por debajo del rango (2.20-5.93 BNN) reportado para poblaciones de murciéla- gos frugívoros neotropicales del continente. Se discuten las posibles causas de esas disminu- ciones en los niveles poblacionales y se presentan comentarios sobre nuestras preocupacio- nes sobre el futuro del estado de conservación de la fauna de murciélagos de la isla.
Key words. Chiroptera. Conservation. Distribution. Saint Martin. Sint Maarten.
Palabras clave. Chiroptera. Conservación. Distribución. Saint Martin. Sint Maarten.
Recibido 26 marzo 2006. Aceptación final 24 abril 2007. 170 Mastozoología Neotropical, 14(2):169-188, Mendoza, 2007 HH Genoways et al. www.sarem.org.ar
INTRODUCTION METHODS AND MATERIALS
The island of St. Martin is the smallest island Study area.—Saint Martin/Sint Maarten (18º05’N, in the world divided between two sovereign 63º12’W) is part of the outer arc of islands of the nations. The northern half of the island is a northern Lesser Antilles. It is a non-volcanic is- part of France (Saint Martin), being adminis- land composed primarily of limestone and igneous diorite. St. Martin occurs on the same submarine tered as a commune of the overseas depart- plateau as Anguilla and St. Barthélemy and formed ment of Guadeloupe. The southern half of the one island with them during the height of Pleis- island is part of the Netherlands (Sint Maarten), tocene glaciations. The island is 15 km long and being administered as part of the Netherlands 14 km wide giving it an area of about 85 sq km. Antilles through Curaçao. Although this sepa- The highest point is Pic du Paradis at 425 m. The ration was hardly perceptible while working month with the highest mean temperature in on the island, it added a layer of administra- Philipsburg is August at 27.9º C and the coolest tive difficulty in conducting a survey of the months are January and February with a mean of chiropteran fauna of the entire island. Over a 24.7º C (Stoffers, 1956). Mean annual rainfall is 1000-1125 mm, but precipitation is very erratic period of three years (2002-2004), we have from year to year and long periods of drought occur. documented eight species of bats from the Only two ravines contained moving water during island of which three are reported from Saint our visits and one of these may have been flowing Martin/Sint Maarten for the first time only because of sewage water being discharged (Monophyllus plethodon, Ardops nichollsi, and into it. Saint Martin/Sint Maarten lies in the Car- Natalus stramineus). ibbean hurricane zone with the late 1990s being The first species of bat recorded from Saint the most active period since the keeping of records Martin/Sint Maarten was based on individuals (1851) with five named hurricanes—Luis, 1995; referable to Artibeus jamaicensis by Cope Bertha, 1996; Georges, 1998; Jose and Lenny, 1999 (Caribbean Hurricane Network, 2005). The most (1889). Subsequently, Husson (1960) in a re- recent storm to hit the island was Debby in 2000. view of the mammals of the Netherlands The most active period for storms for the island is Antilles added three species of bats to the fauna the first week of September (Caribbean Hurricane (Brachyphylla cavernarum, Molossus Network, 2005). molossus, and Tadarida brasiliensis) as well During the early colonial period, the island was as presenting additional information on A. under intense agricultural development with sugar jamaicensis. Finally, Koopman (1968) added cane grown almost to the tops of the hills. Today the greater fishing bat Noctilio leporinus to agriculture has been reduced to a few subsistence the fauna of the island and commented on the areas and pastures. Because of this earlier cultiva- tion, the island is covered with secondary vegeta- other four species known from the island. tion characterized by no definite structure and wide Husson (1960) also reported a single individual variation in composition. The few areas of original of the bat Myotis nigricans nesopolus; how- vegetation are located on the high hills and peaks ever, our re-examination of this specimen in and these are coming under assault today from light of 45 years of additional knowledge of development. The primary original vegetation was the small Myotis in the Lesser Antilles and probably semi-evergreen seasonal forest (Stoffers, northern South America has led us to exclude 1956). At present, the island is under intense de- this species from the fauna of Saint Martin/ velopment pressure to expand tourism and to pro- Sint Maarten. Here, we present information vide space for residential and investment housing and condominiums. about eight species that comprise the current Collecting methods.—Our mist-netting on Saint bat fauna occurring on Saint Martin/Sint Martin/Sint Maarten occurred during three peri- Maarten based on our field research and study ods—12 to 17 July 2002, 31 net nights; 20 to 23 of specimens housed in museum collections. May 2003, 20 net nights; and 14 to 19 May 2004, Based on our work, we present some concerns 35 net nights (86 net nights total). Mist-netting for about the future conservation status of the bats was conducted in a variety of habitats, includ- chiropteran fauna of the island. ing ravines, plantations, covered flyways, trails, and BATS OF ST. MARTIN, NORTHERN LESSER ANTILLES 171 access roads. Due to the season and the dry nature SPECIES ACCOUNTS of the remaining forest on this island, very few locations had any appreciable fruit in evidence. Noctilio leporinus mastivus (Vahl, 1797) Five to eight mist-nets of varying lengths were erected at each location and frequently were situ- Greater Fishing Bat ated diagonally across a trail or ravine at 20 to Specimens examined (2).—SAINT MARTIN: 100 m intervals and monitored for four to six hours Grotte du Puits des Terres Basses, 0.25 km S, 1 depending on bat activity and weather. km W Mornes Rouges, 18º03’48.3"N, Caves and rock faces were surveyed for roosting 63º08’13.3"W, 33 m, 2 (TTU). bats. Two active roosts were found—Grotte du Specimens captured/released (5).—SAINT MAR- Puits des Terres Basses (0.25 km S, 1 km W TIN: Grotte du Puits des Terres Basses, 0.25 km Mornes Rouges) in the lowlands of the western S, 1 km W Mornes Rouges, 18º03’48.3"N, portion of French St. Martin and a cave complex 63º08’13.3"W, 33 m, 5. in the limestone outcrop (Billy Folly) at the east- Additional record.—UNKNOWN: no specific lo- ern edge of Simpson Bay in the southwestern part cality (Koopman, 1968). of Dutch Sint Maarten. Grotte du Puits was visited in 2002 and 2004, and Billy Folly was visited all The greater fishing bat was known previ- three years. A third large cave, Devil’s Hole, was ously from St. Martin based on a single indi- described by Hummelinck (1979). The fate of this vidual deposited in the Academy of Natural cave is discussed in detail in the species account Sciences of Philadelphia from an unknown for Tadarida. Numerous small caves, rock outcrops, location on the island (Koopman, 1968). Davis and rock shelters around Terres Basses, Cul-de- (1973) assigned circum-Caribbean specimens Sac, and Pic du Paradis were searched for evi- to the subspecies N. l. mastivus, which was dence of bat roosts as time permitted during each originally described from St. Croix, U. S. survey but without effect. Virgin Islands. We have followed this arrange- Measurements.—Mist-netted bats were mea- sured and examined at the end of the evening and ment here. Length of forearm and cranial body mass (g), length of forearm (mm), reproduc- measurements of our male and female speci- tive status, tooth wear, presence of scars, and ex- mens are given in Table 1. These measure- ternal parasites were recorded. Length of forearm ments are within the range of those given by and cranial measurements (mm) were also taken Davis (1973) for a sample from the West from museum specimens using digital calipers and Indies. Our male specimen was larger than the weights were recorded in grams. Measurements female in all measurements recorded. were taken following Hall (1946), except that great- Our two specimens were taken on 17 July est length of skull included the incisors, and length 2002 in Grotte du Puits des Terres Basses. A of forearm is the distance from olecranon process to tip of carpals with the wing in retracted posi- general description of this cave is given by tion. StatView® software package (Sager, 1992) Hummelinck (1979). The limestone outcrops provided standard statistics for each sample and throughout this region are covered with low, paired t-tests were used to test for differences thorny dry scrub forest that is dense in many between groups. areas. The cave occurs at the base of a high Voucher specimens.— A survey of existing col- limestone cliff (15 m). The cave entrance is lection materials in natural history museums about 7 m wide and leads to two rooms in (American Museum of Natural History, New York, which the ceiling is about 8 m high. This cave AMNH; Muséum National d’Histoire Naturelle, was mined for phosphate in the late 19th cen- Paris, France, MNHN; Nationaal Natuurhistorisch Museum, Leiden, The Netherlands, RMNH) yielded tury and evidence of this work is quite notice- an additional 63 specimens of bats from Saint Mar- able. In undisturbed portions of the cave, there tin/Sint Maarten. All voucher specimens from the are deep crevices and rounded solution cavi- 2002 survey were deposited in the research collec- ties occupied by bats. Seven individual fish- tions at the University of Nebraska State Museum ing bats, of which two were preserved and (UNSM) and those from 2003 and 2004 were five were released, were taken from one of the deposited in the mammal collections of the Mu- highest solution pockets in the western cham- seum of Texas Tech University (TTU). ber of the cave in 2002. Temperature near the 172 Mastozoología Neotropical, 14(2):169-188, Mendoza, 2007 HH Genoways et al.
www.sarem.org.ar
upper molars upper
Breadth across Breadth
lary toothrow lary Length of maxil- of Length
t Martin, French West Indies.
Mastoid breadth Mastoid
constriction
Postorbital
breadth Zygomatic Table 1 Noctilio leporinus mastivus
Monophyllus plethodon luciae
Ardops nichollsi montserratensis
length
Brachyphylla cavernarum
Condylobasal
of skull of
Greatest length Greatest forearm
87.385.1 29.0 26.3 24.6 23.5 20.1 18.8 7.5 7.0 19.3 16.7 10.3 10.1 12.8 12.1 51.847.3 24.5 22.8 21.4 19.7 15.9 15.1 6.3 5.8 13.0 12.1 8.0 7.4 10.7 9.9 43.7 22.7 21.6 9.7 4.6 9.5 8.2 5.1 Length of Length NMeanRange SE± 55555555 NMeanRange (65.2-67.7) SE± 55555555 66.6 ±0.53 (32.0-33.0) (63.8-67.6) 32.4 65.5 (29.5-29.9) ±0.17 ±0.67 (31.0-32.1) (17.2-18.2) 29.3 31.5 (27.4-28.8) ±0.21 (6.5-6.9) ±0.24 (17.1-17.9) (14.5-15.7) 28.2 17.6 ±0.19 ±0.23 (6.3-6.7) (11.0-11.6) (11.3-12.4) (14.8-15.0) 17.4 ±0.07 6.6 ±0.13 (10.2-11.3) (11.4-12.0) ±0.20 ±0.07 6.5 15.1 ±0.11 ±0.04 11.2 14.8 ±0.21 ±0.19 11.8 10.8 ±0.10 11.6 NMeanRange 55555555 (41.2-43.5) 42.7 (23.5-24.2) 23.8 (20.5-21.6) (10.2-10.6) 21.3 (4.7-4.9) (9.9-10.2) 10.4 (7.9-8.3) 4.8 (5.4-5.9) 10.0 8.1 5.6 Statistics, catalog numbers, and sex TTU 101839, Male TTU 101838, Female TTU 101868, Female TTU 101846, Female Males Females Males SE± UNSM 27997, Female ±0.40 ±0.10 ±0.20 ±0.10 ±0.03 ±0.10 ±0.10 ±0.10 Length of forearm and seven cranial measurements (in mm) for species bats from Sint Maarten, Netherlands Antilles/Sain BATS OF ST. MARTIN, NORTHERN LESSER ANTILLES 173 (Cont. Table 1) Molossus molossus Artibeus jamaicensis Tadarida brasiliensis antillularum 38.1 15.6 14.5 9.1 3.7 8.9 5.6 6.6 NMeanRange SE± 55555555 NMeanRange (59.4-61.5) SE± 55555555 60.7 ±0.37 (27.6-29.4) (56.1-61.8) 28.5 59.8 (24.6-25.4) ±0.30 ±1.02 (28.0-29.2) (16.5-17.4) 25.1 28.4 (25.0-25.9) ±0.12 (7.0-7.5) ±0.23 (16.2-16.9) (14.3-15.0) 25.4 16.9 ±0.19 ±0.18 (7.0-7.3) (9.6-10.0) (14.2-14.8) (12.2-12.9) 16.6 ±0.10 7.2 ±0.16 (9.5-10.2) (12.0-12.7) ±0.15 ±0.05 7.1 14.6 ±0.08 ±0.11 14.6 9.9 ±0.12 ±0.12 12.7 9.9 ±0.12 12.5 NMeanRange SE± 55555555 NMeanRange (37.2-39.8) SE± 55555555 38.6 ±0.45 (16.4-17.1) (38.2-39.3) 16.8 38.6 (14.4-15.0) ±0.11 ±0.19 (16.3-16.7) (10.2-10.8) 14.8 16.5 (14.4-14.5) ±0.11 (3.3-3.4) ±0.07 (10.0-10.3) (9.6-10.0) 14.5 10.4 ±0.11 ±0.02 (3.3-3.4) (5.6-6.1) (9.5-9.7) 10.2 ±0.02 3.3 ±0.07 (7.3-7.5) (5.6-5.9) ±0.08 ±0.0.2 3.3 9.7 (6.7-7.4) ±0.08 ±0.03 9.6 5.9 ±0.04 ±0.06 5.7 7.4 ±0.13 7.2 NMeanRange SE± 55555555 (38.2-38.9) 38.4 ±0.13 (15.9-16.2) 16.0 (14.5-15.0) ±0.13 (9.2-9.5) 14.8 ±0.09 (3.6-3.7) 9.3 (8.7-9.0) ±0.02 (5.4-5.7) ±0.05 3.7 (6.5-6.8) ±0.05 8.9 ±0.06 5.6 ±0.06 6.6 Males Females Males Females Males TTU 101848, Female 174 Mastozoología Neotropical, 14(2):169-188, Mendoza, 2007 HH Genoways et al. www.sarem.org.ar roosting fishing bats was 29º C and the rela- (1967) in placing all populations of this spe- tive humidity was 76%. Other bats roosting in cies from St. Lucia northward in the Lesser this cave in 2002 and 2004 were A. jamaicensis Antilles in the subspecies M. p. luciae, but we and B. cavernarum. In 2002, we spotlighted recognize that the variation represented among the two saline ponds at Little Bay for the pres- samples in the northern Lesser Antilles call ence of foraging N. leporinus without effect. for a revaluation of geographic variation now Of four females captured two were lactating that larger samples are available. The single and two did not evince any gross reproductive female was larger than the mean of the five activity. Testes lengths of three males were 5, males in three measurements (length of the 6, and 8. The four females averaged 58.4 (55.8- forearm, condylobasal length, and length of 61.6) in weight, whereas the three males maxillary toothrow). weighed 71.9, 74.5, and 78.1. Apart from the single male captured near Middle Ground on 14 July 2002, all other Monophyllus plethodon luciae Miller, 1902 specimens were taken at Billy Folly Cave. At Insular Long-tongued Bat Billy Folly Cave, 11 of the 12 specimens taken during visits in July 2002 and May 2003 were Specimens examined (13).—SINT MAARTEN: males. The single female was captured on 15 Billy Folly Cave, 18º01’48"N, 63º05’26"W, 104 July 2002. This segregation of sexes probably m, 12 (7 TTU, 5 UNSM); Romeo’s Drive, Middle indicates that this species forms maternity Region, 18º02’50"N, 63º02’35"W, 74 m, 1 colonies for reproductive females and their (UNSM). young. The 2002 capture of a single male near The single male taken from Middle Ground Middle Ground represents the first record of was netted within a grove of genip trees the long-tongued bat from Sint Maarten/Saint (Melicoecus bijugatus). This locality will be Martin. The species is known from the nearby fully described in the species account for islands of Anguilla (Schwartz and Jones, 1967; Artibeus jamaicensis. The cave complex near Koopman, 1968), St. Kitts (Pedersen et al., Billy Folly is situated within a large limestone 2005), and Nevis (Pedersen et al., 2003) so its outcrop that overlooks both Simpson and Lay presence on the island was not unexpected. bays. This buttress is honeycombed with nu- The species usually frequents dry habitats merous caves and solution pockets, none of where caves are available for daytime roosts. which are readily accessible due to their loca- Forearm and cranial measurements of 5 males tion within a cliff face or are protected by and one female are given in Table 1. These dense thickets of thorny scrub. Bats were lo- measurements when compared with a sample cated in an interconnected pair of caves on from Guadeloupe (Baker et al., 1978) shows the western rim of the buttress. The larger, that the lower end of the range of measure- southern-most chamber was well lighted with ments matches closely between the samples, large openings to the outside through the ceil- but individuals at the upper end of the range ing and western wall and was littered with bat of measurements are larger on Sint Maarten/ guano and bones of goats and iguanas. This Saint Martin for all measurements except for cavity is extensive with numerous side cham- condylobasal length and mastoid breadth. bers that penetrate horizontally back into the Single individuals from Nevis (Pedersen et al., center of the buttress. Bats were found roost- 2003) and Montserrat (Pedersen et al., 1996) ing in the second, northern chamber in both fall below the range of variation of the sample 2002 and 2003. This second chamber was from Sint Maarten/Saint Martin for length of accessible only by a very narrow vertical chim- forearm and greatest length of skull for both ney that descended down into a room of ap- individuals and for mastoid breadth and length proximately 50 cubic meters. The room was of maxillary toothrow for the female from entirely dark with a relative humidity of 89% Montserrat. We follow Schwartz and Jones and a temperature of 29º C while the outside BATS OF ST. MARTIN, NORTHERN LESSER ANTILLES 175
temperature was 32º C. We noted approxi- Martin. Males in our sample were significantly mately 25 bats in 2002 and only 12 in 2003. larger than females in greatest length of skull Seven males collected on 21 May 2003 had and condylobasal length at the P £ 0.05 and P testes lengths that averaged 3.7 (3-5), whereas £ 0.01 levels, respectively. Males also aver- five taken on 15 July 2002 had mean testes aged larger than females in all other measure- lengths of 4.5 (3.5-5). The 12 males weighed ments studied. an average of 16.1 (14.5-18.1). Our single We found Antillean cave bats in only one female revealed no gross reproductive activity cave, Grotte du Puits des Terres Basses, which and weighed 17 when captured. is a site known to earlier researchers (Husson, 1960). During our visit in 2002, Brachyphylla Brachyphylla cavernarum cavernarum were found in both chambers of the cave, but Gray, 1834 in 2004, the maternity colony was confined to Antillean Cave Bat the smaller eastern chamber. In contrast to the dry well, lighted western chamber, the eastern Specimens examined (51).—SAINT MARTIN: chamber was darker, more humid, and had a Grotte du Puits des Terres Basses, 0.25 km S, 1 damp floor covered in guano. Both chambers km W Mornes Rouges, 18º03’48.3"N, 63º08’13.3"W, 33 m, 7 (TTU); upper Loteríe Farm, showed evidence of human disturbance; long 0.5 km S, 1 km E Rambaud, 18º04’41.4"N, wooden poles capable of reaching the roost- 63º03’22.8"W, 75 m, 8 (3 MNHN, 5 TTU); north- ing bats were left leaning against the walls of eastern slope Pic du Paradis, 18º04’43.4"N, both chambers in the vicinity of the roosting bats. 63º02’54.1"W, 410 m, 1 (TTU). SINT MAARTEN: We are not certain of the conditions under Lowlands, 35 (AMNH). which others took individuals at Lowlands, but Specimens captured/released (10).— SAINT circumstantial evidence suggests that they were MARTIN: Grotte du Puits des Terres Basses, 0.25 taken from a cave near the south end of Juliana km S, 1 km W Mornes Rouges, 18º03’48.3"N, Airport. The original vegetation in this area 63º08’13.3"W, 33 m, 10. Additional records.— SINT MAARTEN: small was probably as around Grotte du Puits des cave at Billy Folly (Rojer, 1997: 8); Terre Basse Terres Basses, being low, thorny, dry scrub (Timm and Genoways, 2003). forest, but the area has become so developed Husson (1960) first reported the Antillean that much of the original vegetation is now cave bat from this island based on four speci- gone. This sample includes 17 males and 18 mens from Grotte du Puits des Terres Basses females suggesting that only selected individu- on the French side of the island. The species als were preserved, but it does indicate that also has been reported from the island by pregnant females and adult males were taken Koopman (1968) and Swanepoel and together at this site. Genoways (1978) who reported specimens Rojer (1997) reports and presents a photo- from Lowlands on the Dutch side of the is- graph of a colony of around 300 individuals land. Our work has added material from two of B. cavernarum “in a small cave at Billy additional localities both on the French side Folly.” We explored several small caves of the island. Swanepoel and Genoways (1978) around Billy Folly point during our field stud- assigned members of this species from St. ies in the summers of 2002, 2003, and 2004. Croix southward to St. Vincent to the nomi- The only species of bat that we found occupy- nate subspecies and we have followed that ing these caves was M. plethodon and we never arrangement here. Table 1 presents length of found them in numbers exceeding 25 individu- forearm and cranial measurement for 5 males als. We did expend considerable time sifting and 5 females from St. Martin. Not surpris- through guano and dirt to identify bat remains ingly, the measurements of these specimens but found little more than the fragmented skulls closely match the values for sample 19 in the of 2 or 3 B. cavernarum. study by Swanepoel and Genoways (1978), We also captured foraging individuals on which was composed of specimens from St. two occasions on the slopes of Pic du Paradis. 176 Mastozoología Neotropical, 14(2):169-188, Mendoza, 2007 HH Genoways et al. www.sarem.org.ar
We captured a single adult male along the were captured that were lactating and one re- northeastern slope on the night of 14 May vealed no gross reproductive activity. Six 2004. This individual was taken in a mist net males taken during this period had a mean set across the access road to the top of Pic du testes length of 6.7 (5.5-9.5). Earlier, Husson Paradis on an unseasonably cold night (18.5 (1960) noted that hairless young were found Cº). This area was windswept and the vegeta- in mid-July in Grotte du Puits des Terres tion evinced some wind shaping (“krumholz” Basses. It appears that B. cavernarum has a effect). On the nights of 15 and 16 July 2002, reproductive cycle on St. Martin that includes eight B. cavernarum were netted along the fertilization in February and March, pregnancy western slope of Pic du Paradis in the upper in March, April, and May, and parturition reaches of what is known as Loteríe Farm. extending at least from mid-May to mid-July. Loteríe Farm can best be described as a This is less synchrony than we anticipated in refuge protected by its owner in an attempt to the reproductive cycle of this bat and there is preserve the remaining native vegetation on always the possibility that there are two closely St. Martin and, where possible, the owner is spaced periods of parturition occurring in this trying to restore native vegetation with an cave bat (Wilson, 1979). More data will be aggressive replanting/reforestation effort. This needed to resolve this issue. was the only area that we observed on the Seven males captured in May and July had island that provided good habitat for bats and a mean weight of 48.7 (46.0-52.6) and six other fauna. The deep valley has a small, sea- nonpregnant females taken at these times sonal running stream and several small weighed 48.0 (40.5-53.9) on average. catchement pools of water. The flora in this area varied from evergreen to semi-evergreen Ardops nichollsi montserratensis seasonal forest (Rojer, 1997; Stoffers, 1956). (Thomas, 1894) Antillean cave bats were taken in nets set Lesser Antillean Tree Bat across one of the numerous foot trails on the valley floor and across pools of water. As is Specimens examined (2).—SAINT MARTIN: typical of the species, large mobs of animals lower Loteríe Farm, 0.5 km S, 0.5 km E Rambaud, would forage from tree to tree moving up and 18º04’40.6"N, 63º03’45.9"W, 59 m, 1 (TTU); then down the valley. upper Loteríe Farm, 0.5 km S, 1 km E Rambaud, 18º04’41.4"N, 63º03’22.8"W, 75 m, 1 (TTU). Of eight females captured on 30 March 1926 that were examined for reproductive activity, The discovery of the Lesser Antillean tree seven were pregnant with embryos averaging bat is important because Ardops, despite the 17.4 (13-20) in crown-rump length and one fact that it has been found on several islands evinced no reproductive activity. Four males of the inner arc of the northern Lesser Antillean from this date had testes that averaged 6.0 (5- islands, such as Montserrat (Pedersen et al., 8) in length. On 16 May 2004, three lactating 1996; Jones and Baker, 1979), Nevis (Pedersen females were taken from a maturity colony in et al., 2003), St. Kitts (Pedersen et al., 2005), Grotte du Puits des Terres Basses. Associated and St. Eustatius (Jones and Schwartz, 1967), with these females was a newborn male that has not been reported from any islands in the weighed 14.7 and with a forearm of 36.4. This outer arc, including Antigua (Pedersen et al., individual lacked hair over all of its body. Two 2006), Barbuda, Saint Barthélemy, and slightly older young bats had hair visible on Anguilla. Length of forearm and cranial mea- the dorsum but none on the venter. These two surements of the two females that were cap- males weighed 18.0 and 15.8 and had fore- tured are given in Table 1. These measure- arms of 41.1 and 40.5. A male taken on the ments are slightly smaller than those of a fe- northeastern slope of Pic du Paradis on 15 male from Montserrat reported by Pedersen et May 2004 had testes that measured 7 in length. al. (1996), but closely match those of a female Between 15 and 17 July 2002, two females reported from Nevis by Pedersen et al. (2003). BATS OF ST. MARTIN, NORTHERN LESSER ANTILLES 177
We have tentatively assigned our specimens Romeo’s Drive, Middle Region, 18º02’50"N, from Saint Martin to A. n. montserratensis on 63º02’35"W, 74 m, 9 (3 TTU, 6 UNSM). geographic grounds (Jones and Schwartz, Specimens captured/released (27).— SAINT 1967) and the matching of these measurements, MARTIN: Grotte du Puits des Terres Basses, 0.25 km S, 1 km W Mornes Rouges, 6; lower Loteríe but we recognize a new revision of this genus Farm, 0.5 km S, 0.5 km E Rambaud, 17; upper would be appropriate. Loteríe Farm, 0.5 km S, 1 km E Rambaud, 4. It is our belief that the Lesser Antillean tree Additional records.— SAINT MARTIN: Deux bat occupies a very limited area on Saint Frères (Rojer, 1997: 8). SINT MAARTEN: Martin that may cover as little as 1.5 square Belvédère (Rojer, 1997: 8); St. Peter (Husson, kilometers along the western slope of Pic du 1960); Simpson Bay (Husson, 1960). Paradis (Loteríe Farm). This area probably This was the most common and widespread extends just to the north of the Ravine de species of bat occurring on the island during Loteríe and southward to include the upper our work. The Jamaican fruit-eating bat was slopes of the valley that extends to the south the first species of bat known from St. Martin and east of Colombier. We do not believe that when Cope (1889) described two adult males appropriate habitat extends into Sint Maarten from the island as a new species under the so the species may only be found in Saint name Dermanura eva. Rehn (1901) described Martin. In the area above the great house for the cotypes in the collections of the Academy Loteríe Farm, the habitat was the least altered of Natural Sciences of Philadelphia in more that we observe anywhere on the island, some detail using the name Artibeus eva. Rehn of which we believe may be primary forest (1901) provides the length of forearm of one (Rojer, 1997; Stoffers, 1956). It was in this cotype (ANSP 5784) as 62 and cranial mea- patch of forest that the two Ardops were cap- surements of the second cotype (ANSP 5783) tured on successive nights (16 and 17 May including greatest length of skull 27.6 and 2004) in mist nets that were located within zygomatic breadth 17.0. These measurements 100 m of each other along the trail in the fall within the range of variation for our speci- bottom of the ravine. This suggests that a fa- mens from the island (Table 1) and support vored fruit tree or possibly a roost was lo- the placement of Dermanura eva as a junior cated nearby. Both specimens examined were synonym of Artibeus jamaicensis as was first females and weighed 24.0 and 19.6, respec- done by Andersen (1908) and was continued tively. Neither revealed gross reproductive by Hall (1981). Subsequently, Husson (1960) activity. and Koopman (1968) have reported additional specimens of this species from the island. Artibeus jamaicensis jamaicensis Length of forearm and cranial measurements Leach 1821 for 5 males and 5 females are given in Table Jamaican Fruit-eating Bat 1. These measurements reveal no significant secondary sexual differences between individu- Specimens examined (68).—SAINT MARTIN: als in this sample. Genoways et al. (2001) have Grotte du Puits des Terres Basses, 0.25 km S, 1 reviewed morphological variation in Antillean km W Mornes Rouges, 10 (TTU); lower Loteríe populations of this species. Our sample from Farm, 0.5 km S, 0.5 km E Rambaud, 18º04’40.6"N, 63º03’45.9"W, 59 m, 6 (2 MNHN, 4 TTU); upper Sint Maarten/Saint Martin (Table 1) most Loteríe Farm, 0.5 km S, 1 km E Rambaud, 14 closely matches the measurements of other (TTU); northeastern slope Pic du Paradis, samples of A. j. jamaicensis; in fact, the mean 18º04’43.4"N, 63º02’54.1"W, 410 m, 6 (TTU); SW values of our sample are almost an exact match slope Pic du Paradis, 2 (TTU). SINT MAARTEN: for the means of samples from Puerto Rico Devil’s Hole, Simpson Bay, 4 (AMNH); Mary’s and Dominica (Genoways et al., 2001). Fancy, 0.5 km S Cul de Sac, 18º02’29"N, A maternity colony of Jamaican fruit-eating 63º03’47"W, 26 m, 2 (1 TTU, 1 UNSM); Low- bats was discovered in the eastern room of lands, 4 (AMNH); Philipsburg, 11 (AMNH); Grotte du Puits des Terres Basses where we 178 Mastozoología Neotropical, 14(2):169-188, Mendoza, 2007 HH Genoways et al. www.sarem.org.ar also found the greater fishing bats. Directly area. The fact that we collected nine individu- beneath the roosting bats, the floor of this well- als of the Jamaican fruit-eating bat in two years lit space was covered with almond seeds of work in this region is a tribute to the ability (Terminalia cattapa) and mango pits of this species to tolerate human disturbance. (Mangifera indica). Among the females Certainly, Artibeus feeds on genip fruit, which present, some had not given birth while others may draw the bats into this orchard, but the were carrying newborn on 17 July 2002. Ja- trees also provide a sheltered corridor between maican fruit-eating bats were roosting in the Middle Region and the nearby lowlands of the same general area as the fishing bats, but in a eastern part of the French side of the island. lower part of the ceiling and thus were more Mary’s Fancy is an idiosyncratic semi-aban- easily disturbed. We examined four specimens doned cluster of cottages, trails, a swimming that were taken from the Devil’s Hole cave pool, stream, and a mature collection of large near Simpson Bay (Hummelinck, 1979), but trees and flowering plants including almonds, this valuable cave no longer exists (Rojer, torchwood (Fagara trifoliate), Barbados cherry 1997; see species account for Tadarida). (Malpighia puncicifolia), coconut palms (Co- Among the specimens of this species preserved cos nucifera), banana, mangrove (Rhizophora in the American Museum of Natural History mangle and Avicennia nitida), and other small is a non-volant young male from Devil’s Hole, native trees and shrubs. It exists as an oasis which would indicate that Jamaican fruit-eat- for wildlife amidst the urban clutter of Cul- ing bats reproduced in this cave. The condi- du-Sac. However, only two Jamaican fruit- tions under which Artibeus from Lowlands and eating bats were captured here in two nights Philipsburg were collected are unknown. of netting in 2002 and 2003 a result probably The remaining specimens taken during our due to heavy road traffic nearby, little fruit on research were netted at night in foraging ar- the trees, and dense vegetation, which denied eas. The species seemed particularly abundant the bats access to any available understory in the area of Pic de Paradis and Loteríe Farm. fruits. The native vegetation was less disturbed in An adult female taken on 26 March 1926 these areas than elsewhere on the island, with showed no signs of reproductive activity as native fruit trees and bushes present as well as did a female taken on 30 March 1926, but a a few introduced fruit trees such as mango second female from this date was lactating. and banana (Musa). At Loteríe Farm, fruit bats All seven of the adult females taken on 13 were taken at lower elevations where the veg- April 1926 in Philipsburg were lactating. Three etation was more open and non-native fruit subadult individuals from these dates are pre- trees were more abundant. However, individu- served in collections. Of the five females taken als also were captured high on the northeast- on 15-16 May 2004, one was lactating and the ern slope of the peak as they flew along an other four were pregnant with single embryos access road on Pic du Paradise. that averaged 26.3 (23-32) in crown-rump The specimens from Middle Ground were length. Eight females were captured during our taken in nets set within a small grove of genip work between 13 and 17 July 2002. Of these trees below Romeo Drive in a ravine called 8 females, four were pregnant, with two hav- Cherry Gut by the local residents. Cherry Gut, ing near-term embryos that measured 38 and which was criss-crossed by foot paths and 40 in crown-rump length, whereas two of the contained accumulations of human refuse in- other females evinced no gross reproductive dicating that it was being used as a landfill activity and two were lactating. One of the site, was the most highly disturbed habitat that lactating females from Grotte du Puits des we worked on the entire island. Low scrub Terres Basses on 17 July 2002 was accompa- and weedy plants covered the flanks of the nied by a young pup that had a forearm length ravine, and over-browsing by goats prevented of 39 as compared with mother’s 62. A nearly all but the mature genips from growing in this naked juvenile male taken on 13 April 1926 BATS OF ST. MARTIN, NORTHERN LESSER ANTILLES 179
had a forearm length of 35.5, whereas a young crown-rump length weighed 46.7, 46.8, 49.1, male from this date was covered in juvenile and 54.4. Twelve July-taken adult males pelage and possessed a forearm that was 54.2 weighed an average of 41.2 (36.3-46.7), long. Another young male taken from the whereas two non-pregnant adult females taken Devil’s Hole on 26 March 1926 was covered during this time period weighed 36.0 and 42.0 in juvenile pelage, but had a forearm that was and a lactating female weighed 43.4. Four only 47.1 long. females with near-term embryos when captured Adult males had testes lengths of 6 and 8 on between 14 to 17 July weighed 55.5, 57.0, 30 March 1926 and 7 on 13 April 1926. Five 57.2, and 57.6. adult males taken from Pic du Paradis and A lactating adult female (AMNH 72482) Loteríe Farm, Saint Martin, on 15-16 May captured on 13 April revealed evidence of 2004 had testes that had a mean length of 6.5 alopecia on the top and sides of its head as (6-8), whereas four adult males collected on well as on the chin and extending onto the 21-22 May 2003 from Mary’s Fancy and upper chest area. An adult female (AMNH Middle Region, Sint Maarten, had testes that 72261; length of forearm = 59.3) taken at averaged 8.2 (7-10). Devil’s Hole on 26 March had heavily worn The Jamaican fruit-eating bat is known to dentition, which appeared to be the result of follow a bimodal polyestrous reproductive advanced age of the individual. An adult male cycle in Central America (Wilson, 1979) and taken at Middle Region (TTU 102081) was Jamaica (Genoways et al., 2005). On Jamaica missing the tip of its upper left canine tooth. pregnant females were present in January, The tooth was either broken or had unusually February, and March and in July and August heavy wear, and evidently was infected with and parturition was expected in March-April the abscess draining through the external side and early May and July, August, and Septem- of the alveolus just lateral to the nares. ber (Genoways et al., 2005). The reproductive The presence of the M3/m3 is known to vary data for this species on Sint Maarten/Saint in populations of Artibeus jamaicensis Martin matches the bimodal pattern in data (Genoways et al., 2001). In 10 specimens that for March-April and July reproduction, but we examined from Sint Maarten/Saint Martin, finding pregnant females in mid-May does not the upper M3 was absent in all individuals match the bimodal pattern. Having four of five and the lower m3 was present in seven indi- females pregnant at this time is surprising. viduals, absent in both lower jaws of one in- Given the size of the embryos, they were not dividual, and was present in the right dentary near-term, but certainly would have been born and absent in the left dentary of two individu- before July. This may indicate that the bimo- als. The upper M3 being absent matches other dal polyestrous cycle is not as highly synchro- populations of A. j. jamaicensis, but the per- nous on St. Martin as it is elsewhere in the centage of individuals with the lower m3 geographic range of the species. Alternatively, present is below the 90% or higher levels in this could represent a breakdown in the bimo- other samples of this taxon. dal polyestrous reproductive cycle with females producing as many young in a year as pos- Natalus stramineus stramineus Gray 1838 sible in response to low population levels on Lesser Antillean Funnel-eared Bat the island or resource availability (Fleming, 1971). This observation deserves further study Specimens examined (0). to verify its occurrence and to clarify its sig- Additional record.—SINT MAARTEN: Billy nificance. Folly Cave, 18º01’48"N, 63º05’26"W, 104 m. Nine males taken in mid-May had a mean During the first visit to the Billy Folly Cave weight of 41.0 (37.9-45.5). A lactating female system in 2002, one member of our party captured on 16 May weighed 40.4 and four (Pedersen) observed and attempted to collect females with mid-term embryos based on the skeletal remains of a small bat in the south- 180 Mastozoología Neotropical, 14(2):169-188, Mendoza, 2007 HH Genoways et al. www.sarem.org.ar ernmost chamber. The partial cranium, jaw, Airmouse Cave), could have had a negative and some post-cranial elements were located impact on the status of this species, which is on a protected small rock shelf about a meter dependent on dark, moist caves as day roosts above the floor of the cave. Although these (Genoways et al., 2001). fragments proved to be too fragile for collec- tion, Pedersen was able to identify them as Tadarida brasiliensis antillularum Natalus stramineus based on the distinctive (Miller, 1902) shape of the cranium and dentary. This por- Brazilian Free-tailed Bat tion of the cave was relatively open and a Specimens examined (6).—SAINT MARTIN: headlight was not necessary to see the remains. lower Loteríe Farm, 0.5 km S, 0.5 km E Rambaud, This is not the typical cave situation in which 18º04’40.6"N, 63º03’45.9"W, 59 m, 4 (TTU). SINT we have observed colonies of Natalus on other MAARTEN: Mary’s Fancy, 0.5 km S Cul de Sac, islands in the northern Lesser Antilles. 18º02’29"N, 63º03’47"W, 26 m, 2 (UNSM). Dávalos (2005) and Tejedor et al. (2005) Additional record.—SINT MAARTEN: Bat Cave, presented evidence that unrecognized taxa of Devil’s Hole, Simpson Bay (Husson, 1960; Natalus exist within currently named popula- Hummelinck, 1979). tions in the West Indies. Goodwin (1959), in This free-tailed bat was first reported from his revision of members of the subgenus Sint Maarten by Husson (1960) based on 24 Natalus, restricted the type locality of Natalus female specimens collected in the Bat Cave at stramineus to Antigua rather than Lagoa Santa, Devil’s Hole by Hummelinck (see Minas Gerais, Brazil, as was done by earlier Hummelinck, 1979). Although this large col- authors. This decision later was reconfirmed lection from a single place on Sint Maarten by Handley and Gardner (1990) and Tejedor would seem to indicate that Tadarida was (2006). Tejedor (2006) restricted N. stramineus abundant on the island, we found the species to the northern Lesser Antilles, ranging from to be quite rare being represented by only 6 Dominica to Anguilla, and in a canonical analy- individuals from two localities, one located sis of external and cranial characters, demon- on each side of the island. Length of forearm strated that “the sample from Dominica is and cranial measurements for 5 males and 1 morphometrically distinct from that of the female are reported in Table 1. The length of remaining islands.” Although Tejedor (2006) forearm of specimens from Sint Maarten/Saint did not use it, the name N. s. dominicensis Martin appears to average larger than those of Shamel (1928) is available for the population a sample from the island of Dominica on Dominica, leaving the nominate subspecies, (Genoways et al., 2001), but the cranial mea- N. s. stramineus, as the appropriate name to surements seem to match closely between these apply to the other populations in the northern samples. We continue to follow Schwartz Lesser Antilles, including those on Sint (1955) in assigning all Lesser Antillean popu- Maarten/Saint Martin. lation of this species to the subspecies T. b. We are not able to state with certainty antillularum until a comprehensive study of whether populations of Natalus are extant on geographic variation in Antillean populations the island. We believe that the remains ob- is undertaken. Owen et al. (1990) presented served in Billy Folly Cave had been deposited evidence of a close relationship between T. b. there in the last 50 years, but with the envi- cynocephala from the southeastern United ronmental changes that have occurred on Sint States and Antillean populations of T. Maarten/Saint Martin over this period of time, brasiliensis. Further data are needed to fully this species could have been lost from the explore the taxonomic and biogeographic im- fauna. Our work revealed a limited number of plications if such a relationship exists. caves on the island and Hummelinck (1979) The measurements of our female specimen described only two caves on the island. The were less than the mean of the sample of males loss of the largest cave, Devil’s Hole (or for four measurements—length of forearm, BATS OF ST. MARTIN, NORTHERN LESSER ANTILLES 181
greatest length of skull, condylobasal length, there was no evidence of guano on the floors, and zygomatic breadth, but only in the last apparently the bats did not relocate here when measurement was its value below the range of the main chamber was filled in. The loss of measurements for the males. In the other four this maternity cave due to resort development measurements, the values for the female were must have been a major blow to the popula- the same as the mean for the sample of males. tion of the Brazilian free-tailed bats on the The single most important known site for island and undoubtedly accounts, at least in Brazilian free-tailed bats on the island was the part, for why this free-tailed bat is relatively Bat Cave at Devil’s Hole. This cave is de- rare on St. Martin. scribed in detail by Hummelinck (1979). He During our work in Sint Maarten, we cap- found this natural cave to consist of a single tured two adult male Brazilian free-tailed bats room of about 25 by 15 m. The room was on 13 July 2002 in a 10-m mist net that was accessible because part of the ceiling had set over the abandoned swimming pool at collapsed, but bowl-shaped solution holes were Mary’s Fancy. The pool was located in an evident in the remaining portions of the origi- opening to the sky in this heavily wooded area. nal roof. The cave had been mined in the late The pool was half-filled with one end covered 19th century so its original shape was un- in water hyacinths. Four individuals of Pallas’ known. The floor of the cave was covered in mastiff bats (M. molossus) also were taken over guano and pools of water. Some of these pools the pool. When we returned to this pool in were tidal/brackish because the eastern shore 2003, it was completely covered in water plants of Simpson’s Bay was approximately 100 m and no bats were captured. from the cave. This main chamber contained a On the French side of the island, we cap- nursery colony composed primarily of adult tured four individuals at Loteríe Farm—two female Tadarida brasiliensis. Hummelinck males and a female on 16 May 2004 and one (1979) mentions that the Devil’s Hole had been male on 18 may 2004. They were all taken in “adapted to tourist purposes” just before the nets set in open areas between widely scat- appearance of his publication. Hummelinck tered large trees in the vicinity of the farm (1979) also describes two near-cylindrical buildings. Short grass covered the ground mining shafts in the immediate vicinity of the between the trees. Specimens of M. molossus Bat Cave but these shafts contained no bats. and A. jamaicensis also were taken in this area. We were unable to find Devil’s Hole during The testes lengths of three May-taken males our field research in 2002 and 2003, but we were 2.5, 3.0, and 3.0 and those of two July- continued to try to determine its location dur- taken males were 2.0 and 2.5. The single fe- ing our third season of work in 2004. The male in our sample was taken on 16 May 2004, next to the last day of our work on the island evinced no gross reproductive activity, and we came to an active construction site in the weighed 9.1. The weights of the five males quickly developing housing area above the were, respectively, 9.2, 8.8, 9.1, 9.4, and 9.4. southeast shore of Simpson’s Bay. We inquired of some of the workers at the site about their Molossus molossus molossus (Pallas, 1776) knowledge of the Devil’s Hole. They quickly Pallas’ Mastiff Bat responded that it was here in front of us. According to their account, the main part of Specimens examined (23).—SAINT MARTIN: the cave had been filled-in during the early lower Loteríe Farm, 0.5 km S, 0.5 km E Rambaud, 1990s; however, the two mine shafts still re- 18º04’40.6"N, 63º03’45.9"W, 59 m, 8 (1 MNHN, mained although they were being backfilled 7 TTU); northeastern slope Pic du Paradis, 18º04’43.4"N, 63º02’54.1"W, 410 m, 1 (TTU); as we watched. We were able to quickly ex- southwestern slope Pic du Paradis, 18º04’N, plore them and found that the entrances were 63º03’W, 350 m, 2 (TTU). SINT MAARTEN: about 75% filled with rock rubble and trash. Mary’s Fancy, 0.5 km S Cul de Sac, 18º02’29"N, There were no bats in these mine shafts and 63º03’47"W, 26 m, 6 (2 TTU, 4 UNSM); 182 Mastozoología Neotropical, 14(2):169-188, Mendoza, 2007 HH Genoways et al. www.sarem.org.ar
Philipsburg, 2 (AMNH); Romeo’s Drive, Middle Indies (Gannon et al., 2005; Genoways et al., Region, 18º02’50"N, 63º02’35"W, 74 m, 4 (1 TTU, 2005; Pedersen et al, 2005, 2006). Foraging 3 UNSM). M. molossus were taken in nets along with Specimens captured/released (33).—SAINT T. brasiliensis over the abandoned swimming MARTIN: lower Loteríe Farm, 0.5 km S, 0.5 km pool at Mary’s Fancy and the open grassy area E Rambaud, 18º04’40.6"N, 63º03’45.9"W, 59 m, 33. with isolated single large trees at lower Loteríe Additional record.—SINT MAARTEN: Farm and along with A. jamaicensis in the Philipsburg (Husson, 1960); ruins of “One Hoe” ravine at Romeo’s Drive in Middle Region. [18º03’N, 63º02’W] (Rojer, 1997: 8). Elsewhere, individuals were netted over roads Husson (1960) was the first to report this along the northeastern and southwestern slopes free-tailed bat from Sint Maarten based on 21 of Pic du Paradis, which we believe were used specimens collected by Hummelinck in as protected flyways to move from one side of Philipsburg in 1955. Koopman (1968) reported the island to the other. an additional two specimens from Sint Maarten Of the 10 females taken between 15 and 22 in the collection of the American Museum of May during our work on the island, eight re- Natural History, which undoubtedly were the vealed no gross reproductive activity, whereas two specimens from Philipsburg listed above. single females taken on 16 and 18 May car- This mastiff bat was more abundant and wide- ried embryos that measured, respectively, 7.5 spread on the island during our work than the and 6.0 in crown-rump length. Four males other free-tailed species, T. brasiliensis. taken during this same time period had testes Husson (1962) restricted the type locality of that averaged 3.4 (2-5) long. Three females, M. molossus to the island of Martinique, which prepared as specimens on 13 and 14 July, lead Dolan (1989) to apply the name M. m. revealed that one taken on each date was preg- molossus to this species throughout the Lesser nant carrying a single embryo that measured, Antilles. Length of forearm and cranial mea- respectively, 22 and 9 in crown-rump length, surements for five males and five females from whereas the third female evinced no gross Sint Maarten/Saint Martin are given in Table 1. reproductive activity. Four males taken during These measurements compare favorably with this same time period had testes that averaged samples from Antigua (Pedersen et al., 2006), 4.5 (4-5) long. Eleven of 12 mastiff bats cap- Nevis (Pedersen et al., 2003), and St. Kitts tured and released on the night of 15 May (Pedersen et al., 2005). Based on the work of 2002 were females. Ten of the females were these authors, we have assigned our material judged to be pregnant with the eleventh fe- to the nominate subspecies. Males were sig- male being small, weighing only 9.7 and be- nificantly larger than females only for ing judged as non-reproductive. Because condylobasal length at the P £ 0.05. Males Genoways et al. (2005) on Jamaica found fe- averaged larger, although not at a significance males of this species at various stages of re- level equal to or below P £ 0.05, than females production and others reproductively inactive in all other measurements except length of during the same period, they concluded that forearm and postorbital constriction for which the reproductive cycle most closely resembled the sexes had the same mean values. an aseasonal polyestrous pattern (Wilson, All of the specimens of Pallas’ mastiff bat 1979). Our data from St. Martin are not in taken during our work were netted in foraging conflict with this conclusion, but far more data areas. However, Husson (1960) reported that are needed to confirm this conclusion. the first specimens collected from the island The eight non-pregnant females taken dur- were taken by Hummelinck from the attics of ing May weighed on average 12.1 (9.9-16.0), schools and houses, whereas Rojer (1997) whereas the non-pregnant female taken on July reported this species as roosting among the 14 weighed 10.6. In comparison, the 10 preg- ruins at “One Hoe.” These are typical roost nant females that were captured and released sites for this species elsewhere in the West in May weighed on average 13.7 (12.6-14.4). BATS OF ST. MARTIN, NORTHERN LESSER ANTILLES 183
The four males taken in May weighed on of the length of the tibia to the forearm. LaVal average 12.2 (11.2-13.0) and the four males (1973) stated “The ratio of 0.35 [for M. taken in July weighed on average 13.1 (11.9- dominicensis] is exceeded by all other series 13.7). of Neotropical Myotis I have examined. A large majority of the series of all species fall within SPECIES REMOVED FROM FAUNA the range of 0.38 to 0.42, except Myotis larensis, with a ratio of 0.48.” With a tibia Myotis nesopolus Miller, 1900 length of 16.0, the ratio of the length of the tibia to the forearm for the specimen from St. Specimen examined (1).—SINT MAARTEN: Cul Maarten is 0.51. This ratio would certainly de Sac, 1 (RMNH). argue against assigning the specimen from St. Husson (1960) first reported this specimen Maarten to M. dominicensis and would seem from Sint Maarten under the scientific name to align it with M. larensis, which is a species Myotis nigricans nesopolus. Since this publi- described by LaVal (1973) from the dry coastal cation, the taxonomy of the Myotis nigricans areas of northern Venezuela characterized by group has undergone considerable change, small size, but very long third metacarpal, tibia, particularly by LaVal (1973). LaVal (1973) and greatest length of skull in comparison with assigned West Indian bats of this genus to two the forearm. Genoways and Williams (1979) species—Myotis dominicensis from the island demonstrated that M. larensis was best con- of Dominica and Myotis martiniquensis from sidered to be a subspecies of M. nesopolus, a the islands of Martinique and Barbados. LaVal species originally described from Curaçao and (1973) did not comment on this specimen from not treated by LaVal in his study. Table 2 St. Maarten or two specimens from Grenada gives values for these three ratios for seven (Koopman, 1968). Masson and Breuil (1992) taxa of Myotis from Caribbean islands and Car- subsequently have reported a Myotis from ibbean coast of South America. The values of Guadeloupe that presumably would be assigned the specimen from St. Maarten clearly fall with to M. dominicensis, at least on geographic those of the taxa of Myotis nesopolus from grounds. Koopman (1968) initially assigned coastal Venezuela, Curaçao, and Bonaire. the specimens from Grenada to M. n. nigricans Morphologically, the specimen from St. and Genoways et al. (1998) added a third Maarten most closely resembles Myotis specimen for the island and supported nesopolus from two islands just off the north- Koopman’s taxonomic assignment. Therefore, ern coast of Venezuela and the adjacent main- as is currently understood, three species of land. This brings the discussion back to Husson small Myotis occur in the Lesser Antilles and (1960) who stated that “It can’t be said with the species most closely approaching Sint certainty whether the specimen from St. Mar- Maarten is M. dominicensis known from tin really belongs to M. nesopolus; however, Guadeloupe about 240 kilometers to the south. the specimen resembles M. nesopolus more The measurements of the specimen from St. than it does M. nigricans dominicensis.” We Maarten (see below) do more closely resemble are in full agreement with this statement. If those of M. dominicensis and M. nigricans we accept that the specimen from St. Maarten than they do the measurements of the larger- is a representative of M. nesopolus, we have sized M. martiniquensis (LaVal, 1973: 35). a species occurring on the northeastern edge However, length of forearm of the specimen of the Caribbean Sea and two islands just off from St. Maarten at 31.1 is considerably the north coast of South America more than shorter than averages of the former two spe- 900 kilometers to the southwest. This is not a cies at 34.0 and 34.8, respectively. The single pattern of distribution known for any other characteristic that LaVal (1973) found that species of Antillean bat so an explanation of distinguished M. dominicensis was a low ratio this distribution pattern needs to be considered. 184 Mastozoología Neotropical, 14(2):169-188, Mendoza, 2007 HH Genoways et al. www.sarem.org.ar
Table 2 Ratios of the length of third metacarpal, tibia, and greatest length of skull to the length of forearm in seven taxa of Myotis and the specimen of Myotis from Sint Maarten. The taxa of Myotis are from the Lesser Antilles, northern coast of Venezuela, Curaçao, and Bonaire.
Species Length of third metacarpal/ Length of tibia/ Greatest length of skull/ Length of forearm Length of forearm Length of forearm
Myotis dominicensis 0.881 0.35 ± 0.0042 0.381 Myotis martiniquensis1 0.90 0.38 0.39 Myotis nigricans2 0.93 ± 0.006 0.41 ± 0.005 0.40 ± 0.002 Myotis albescens2 0.92 ± 0.014 0.42 ± 0.009 0.41 ±0.011 Myotis keaysi2 0.90 ± 0.007 0.40 ± 0.006 0.38 ± 0.009 Myotis nesopolus larensis2 0.96 ± 0.003 0.48 ± 0.005 0.43 ± 0.005 Myotis nesopolus nesopolus3 0.97 0.48 0.43 Myotis from St. Maarten 0.95 0.51 0.44
1 Calculated using data from LaVal, 1973. 2 Values taken from LaVal, 1973. 3 Values taken from Genoways and Williams, 1979.
We only know from Husson (1960) that the maxillary toothrow, 5.1; breadth across upper specimen was found in an old building in Cul molars, 4.9. de Sac on the Dutch side of the island. It was obtained by F. A. da Costa on 20 September DISCUSSION 1946. In searching for an explanation for this distribution pattern, it is difficult to ignore the Faunal composition.—Data presented herein fact that the Curaçao, Bonaire, and Sint provide records of three species of bats new Maarten are, and have been, members of the to the fauna of the island of Saint Martin/Sint Netherlands Antilles. Also the fact that the Maarten—Monophyllus plethodon, Ardops specimen was collected just following World nichollsi, and Natalus stramineus. With the War II when traffic among the islands was addition of these three species and the omis- high needs to be considered. Our tentative sion of Myotis nigricans nesopolus, the chi- conclusion based on the facts and circumstan- ropteran fauna of the island is composed of tial evidence is that the specimen from Sint eight species. Ecologically, this simple chi- Maarten represents an individual that was ac- ropteran fauna includes representatives of four cidentally transported from Curaçao or Bonaire families—Noctilionidae, Phyllostomidae, to the island. Our work on the island provided Natalidae, and Molossidae—including one no evidence of a resident population of Myotis, piscivore (N. leporinus), one omnivore (B. although such a negative hypothesis can’t be cavernarum), one pollenivore/nectarivore (M. proven. We believe at the present time it is plethodon), two frugivores (A. nichollsi, A. best to remove Myotis from the chiropteran jamaicensis), and three insectivorous species fauna of Sint Maarten pending any evidence (N. stramineus, T. brasiliensis, and M. to the contrary. molossus). External and cranial measurements of the As our knowledge of the bats occurring on male from St. Maarten (RMNH 17107) are as the Lesser Antillean islands north of follows: length of forearm, 31.1; length of third Guadeloupe increases, an interesting biogeo- metacarpal, 29.4; length of tibia, 16.0; great- graphical pattern may be emerging. The chi- est length of skull, 13.6; condylobasal length, ropteran fauna of Nevis (Pedersen et al., 2003) 12.7; zygomatic breadth, 7.6; postorbital con- exactly matches that of Saint Martin/Sint striction, 3.2; mastoid breadth, 6.7; length of Maarten. The faunas of St. Kitts (Pedersen et BATS OF ST. MARTIN, NORTHERN LESSER ANTILLES 185
al., 2005) and Antigua (Pedersen et al., 2006) Pedersen et al. 2005). We use this BNN met- each are known to have seven species of bats. ric conservatively and avoid its use as an es- The only species missing from St. Kitts that is timate of population size per se, rather BNN known from Saint Martin/Sint Maarten is is best used as an approximation of bat activ- Natalus stramineus, and Ardops nichollsi is ity at a particular location. Indeed, Fenton et not present on Antigua. The former is not a al. (1992) showed that phyllostomid bats are surprise given the difficulty of capturing this useful indicators of habitat disruption; they funnel-eared bat, but the latter is more puz- documented a 21% decrease (4.20 > 3.29 zling given the abundance of Antillean tree BNN) in fruit bat capture rates in their com- bats on several adjacent islands. parison of undisturbed with disturbed habitats On Montserrat the fauna contains an addi- in Mexico. Island populations of fruit bats tional two species—Sturnira thomasi and appear more susceptible to habitat disruption Chiroderma improvisum—that do not occur than are mainland populations. Indeed, if con- on any island north of Guadeloupe except for trasts between pre- and post-hurricane surveys the small populations on Montserrat. These are made using comparable data from data seem to indicate that the bat fauna of Montserrat, captures rates decrease by as much Montserrat with its 10 species has a stronger as 60 to 80% after a natural disaster (Pedersen biogeographic relationship with Guadeloupe et al., 1996; Pedersen, 2001). Our observa- than with other islands in the northern Lesser tions of the remaining forest of Saint Martin/ Antilles. The other islands in the northern Sint Maarten and the conditions for bats in Lesser Antilles would appear to share a fauna particular were not very encouraging and sug- that contains at most eight species of bats such gest an anthropogenic ecological disaster. as those present on Nevis and Saint Martin/ In several areas on the island, we encoun- Sint Maarten. We need data from additional tered accumulating piles of garbage and hu- islands to confirm this observation, but many man waste. The long-term concern is that these of these are either small—Saba, Sint Eustatius, areas will lose all native vegetation and will and Saint Barthélemy—or are low lying— be of no more use to the wildlife and more Anguilla and Barbuda. It will be interesting to importantly to humans, including tourists. The learn if these islands share this same fauna of watertable of the island has been reduced to eight species. the point that we observed only two small flow- Conservation.—Although the bat fauna of ing streams and the one through Dutch Cul de Saint Martin/Sint Maarten is not unique, we Sac was so polluted with oil and phosphates must express considerable concern about the that bats could not use it as a water source. future of each species. If all feeding guilds are We noted very few freshwater ponds on the considered, bat capture rates on Sint Maarten/ island and most reservoirs were covered, leav- Saint Martin (1.63 bats per net per night - ing resort and residential swimming pools as BNN) fall well below capture rates reported the only source of freshwater available to bats from mainland populations (2.59-6.65; Fenton and other wildlife. et al., 1992; Findley, 1993) and are among the In conjunction with the desire for expansion lowest noted on islands throughout the Lesser of the tourism industry, there is a major build- Antilles that the authors have surveyed (1.55- ing boom on the island. Residential building 3.75; Pedersen et al., 2005). If only fruit bats of upscale homes, villas, and condominiums are evaluated, capture rates during our three is progressing at a rapid pace, especially on surveys on Sint Maarten/Saint Martin (0.92 the Dutch half of the island. On the French BNN) are again on the low end of those rates side the construction of access roads and pri- reported from adjacent islands in the northern vate drives on the steep slopes of Pic du Lesser Antilles (0.65-2.47), and fall well be- Paradis (425 m) have created great scars on low fruit bat captures per net-night in the the landscape and severely altered the ability Neotropics of Central America (2.20-5.93; of the soil to store water. Private residences 186 Mastozoología Neotropical, 14(2):169-188, Mendoza, 2007 HH Genoways et al. www.sarem.org.ar are found at 410 m on the northeastern slope the Devil’s Hole would have been a far more and up to 350 m on the southwestern slope. appropriate environment for the species than The subsequent loss of native vegetation im- is provided by Billy Folly Cave. We are cer- pacts not only the food resources available to tain that there are other small caves on this bats, but also deprives some species (Ardops island, but they are not the large cave com- nichollsi and Artibeus jamaicensis) of roost sites. plexes that we have found on adjacent islands, The removal of native vegetation has prob- such as Anguilla or Barbuda. ably had an impact on the insect fauna of the The most important thing that can be done island, which was suggested by Steadman et to conserve chiropteran populations on Saint al. (1984) as a possible cause of the loss of Martin/Sint Maarten is to protect all of the two species of insectivorous bats on Antigua. remaining cave resources on the island. The The most striking example of potential reduc- caves and their immediate environs should be tion of an insectivorous species was the low preserved as natural areas so that the impact numbers of the normally over-abundant of human activity and development can be Molossus molossus of which we took only 49 limited in the future. Caves are very highly specimens in our three years of work giving a specialized ecosystems that possess a signifi- capture rate of 0.57 bats per net night (M. cant amount of biological diversity. The caves molossus—Montserrat: 0.98; St. Kitts: 1.14; are certainly very important to the continued St. Eustatius: 1.97; Nevis: 2.41 BNN). The survival of many species of bats on Saint loss of a diversity of food sources in the form Martin/Sint Maarten, but on the other hand, of insects seems the most likely reason for the bats are extremely important to these ecosys- reduction of this species on Saint Martin/Sint tems because they perform the same role for Maarten because their preferred roost sites caves as green plants do for non-cave situa- associated with tile and corrugated steel roofs tions. It is the dropped food items and guano appear to be abundant on the island. of bats that are primary sources of energy Certainly, the heaviest blow to the chi- entering these ecosystems. To maintain a ropteran fauna of the island was the filling of healthy set of cave ecosystems with their rich the Devil’s Hole cave and its two associated biodiversity, it will be extremely important to tunnels. This was the largest known bat cave maintain healthy bat populations on Saint on the island and had reproductive popula- Martin/Sint Maarten (Genoways et al., 2005). tions of at least Tadarida brasiliensis and We don’t wish to paint only a dark picture Artibeus jamaicensis and probably also could of the future of bat conservation on Saint have served Brachyphylla cavernarum. Half Martin/Sint Maarten. There are many dedicated of the species of bats on the island—M. people working in a variety of conservation plethodon, B. cavernarum, N. stramineus, and projects across the island (primarily concern- T. brasiliensis—are obligate cave roosting ing marine ecosystems) and our work would species, meaning that they require caves for have been impossible without their full coop- day roosts and nursery colonies of newborn eration. They labor under severely limited bats. Noctilio leporinus and Artibeus funding with limited personnel. The Dutch jamaicensis are facultative cave species, mean- have an excellent terrestrial conservation plan ing that they will use caves when available for that was developed in the mid-1990s (Rojer, reproduction, day roosts, and to escape harsh 1997), but during our survey work, it was not environmental condition, such as hurricanes. apparent that the plan had been implemented. The few scattered remains of the funnel-eared This plan deserves official support and imple- bat, Natalus stramineus, were found in Billy mentation to help preserve the overall Folly Cave just up slope from the former lo- biodiversity of the island. A real conservation cation of Devil’s Hole. We question the cur- bright spot was Loteríe Farm in French Saint rent presence of this species on island because Martin. The owners of this private property BATS OF ST. MARTIN, NORTHERN LESSER ANTILLES 187 are making significant progress in preserving LITERATURE CITED and restoring the native vegetation of the prop- erty. This is an exciting development because ANDERSEN K. 1908. A monograph of the chiropteran we believe the most important areas for pres- genera Uroderma, Enchisthenes, and Artibeus. Pro- ceedings of the Zoological Society of London 1908: ervation on the island are headwaters of the 204-319. Loterie and Columbier valleys along the west- BAKER RJ, HH GENOWAYS, and JC PATTON. 1978. ern slopes of Pic du Paradis. These valleys Bats of Guadeloupe. Occasional Papers of the Mu- retain the most extensive areas of native and seum, Texas Tech University 50:1-16. CARIBBEAN HURRICANE NETWORK. 2005. St. advanced secondary vegetation remaining on Maarten/St. Martin (TNCM). Accessed November the island. They should have the highest pri- 21 at
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