TEXAS TECH UNIVERSITY Natural Science Research Laboratory Special Publications Museum of Texas Tech University Number 69 29 November 2018 Bats of Saint Lucia, Lesser Antilles Scott C. Pedersen, Gary G. Kwiecinski, Hugh H. Genoways; Roxanne J. Larsen; Peter A. Larsen, CarletonJ. Phillips; ajvd Robert J. Baker Front cover: The bats of Saint Lucia (from upper left to lower right): (row 1) Monophyllusplethodon, Brachypkylla cavernarum, Sturnirapaulsoni; (row 2) Artibeus jamaicensis x schwartzi, Ardops nichollsi, Pteronotus davyi; (row 3) Noctilio leporinus, Molossus molossus, and Tadarida brasiliensis. Photographs by Gary G. Kwiecinski. Special Publications Museum of Texas Tech University Number 69 Bats of Saint Lucia, Lesser Antilles Scott C. Pedersen, GaryG. Kwiecinski, HughH. Genoways, Roxanne J. Larsen, Peter A. Larsen, CarletonJ. Phillips, and Robert J. Baker Layout and Design: Lisa Bradley Cover Design: Gary G. 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Bradley Bats of Saint Lucia, Lesser Antilles Scott C. Pedersen, Gary G. Kwiecinski, Hugh H. Genoways, Roxanne J. Larsen, Peter A. Larsen, Carleton J. Phillips, and Robert J. Baker ISSN 0149-1768 ISBN 1-929330-36-7 ISBN13 978-1-929330-36-2 Museum of Texas Tech University Lubbock, TX 79409-3191 USA (806)742-2442 Bats of Saint Lucia, Lesser Antilles Scott C. Pedersen, Gary G. Kwiecinski; Hugh H. Genoways, Roxanne J. Larsen, Peter A. Larsen, CarletonJ. Phillips; and Robert! Baker Abstract Eight species of bat have been previously recorded from the island of Saint Lu¬ cia: Noctilio leporinus, Monophyllus plethodon, Artibeus jamaicensis, Brachyphylla cavernarum, Ardops nichollsi, Sturnira paulsoni, Molossus molossus, and Tadarida brasiliensis. Herein, we add a ninth species to the fauna—Pteronotus davyi. These nine species represent nine genera from four families: Noctilionidae, Mormoopidae, Phyllostomidae, and Molossidae. This fauna includes four trophic guilds: N. lepori¬ nus (piscivore/insectivore), M. plethodon (nectarivore/pollenivore), A. jamaicensis x schwartzi, B. cavernarum, A. nichollsi, and S. paulsoni (frugivores), and P. davyi, M. molossus, and T. brasiliensis (insectivores). Based on its geographic location, the bat fauna of St. Lucia is enigmatic in regard to species dispersal along the Lesser Antillean archipelago. Natalus stramineus and M. martiniquensis are present on Martinique im¬ mediately to the north, and Micronycteris buriri, P. fuscus, Glossophaga longirostris, and Artibeus lituratus are found on St. Vincent directly to the south, yet these six species are conspicuously absent on St. Lucia. Moreover, the St. Lucian population of Artibeus resides at the northern edge of a fascinating hybrid zone involving three species, A. jamaicensis, A. planirostris, and A. schwartzi. In light of these observations, we provide a framework for analyzing the biogeographical patterns of the regional fauna to see if there is something about St. Lucia that is unique. We posit that a number of complex geological and ecological factors account for the depauperate bat fauna observed on the island as well as the formation of the Artibeus hybrid zone in the southern Lesser Antilles. Key words: biological invasion, Chiroptera, environmental disruption, hybrid swarm, Mammalia, natural dispersal, natural history, propagule, Pteronotus davyi, reproduction, Saint Lucia, species richness Introduction The first report of a bat from St. Lucia was by and Genoways 1978). In this study, we added the ninth Dobson (1878) based on an adult male Molossus molos¬ species to the fauna—Pteronotus davyi. sus in the British Museum (Natural History), which had been purchased from Miss R. Alexander. Subsequently, Perhaps the most remarkable aspect of the bats of seven additional species of bat were collected on St. St. Lucia is that the resident population of Artibeus in¬ Lucia by several individuals, notably: H. S. Branch in cludes hybrids between A. jamaicensis and A schwartzi 1901 (NMNH) and J. L. Peters in 1925; J. Knox Jones, (P. Larsen et al. 2010). Indeed, the island is located Jr., and C. J. Phillips in 1967 (KU); and J. J. Gulledge at the northern edge of an active hybrid zone among in 1971 (AMNH). Their voucher specimens are well multiple species of Artibeus and therefore traditional reported in the literature (Dobson 1878; Miller 1902, taxonomic assignment of species-level status is prob¬ 1913a, 1913b; Allen 1908, 1911; Andersen 1908; lematic. The fauna also includes S. paulsoni, a species Shamel 1931; Jones and Schwartz 1967; Koopman endemic to three southern Lesser Antillean islands, 1968; Jones and Phillips 1970; Jones 1978; Swanepoel and A. nichollsi, endemic to the Lesser Antilles but not 1 2 Special Publications, Museum of Texas Tech University found on the southern islands of Barbados and Grenada. Members of our research team first visited St. Two species—B. cavernarum and M. plethodon—are Lucia 26 to 28 August 1967 when C. J. Phillips and Antillean endemics with geographic ranges extending the late J. K. Jones, Jr., worked two sites in Dauphin from Puerto Rico (Monophyllus appearing only as a and Gros Islet quarters. Subsequently the island was fossil) in the Greater Antilles southward to the islands visited 24 to 26 May 1987, when H. H. Genoways and of St. Vincent and Barbados. Of those species occur¬ C. J. Phillips focused on the collection of Artibeus near ring on St. Lucia, three (Nleporinus, P. davyi, and M. Marigot Bay. We returned to the island to net additional molossus) have entered the Lesser Antilles directly bats on the following dates: 15 to 20 June 2007; 29 from the South American mainland or via Trinidad, July to 3 August 2008; and 13 to 16 March 2009. It whereas the fourth species (T. brasiliensis) entered the is the combination of these data that we report herein islands from either the west via the Yucatan Peninsula to elucidate the composition and relationships of the or Nicaragua, or from Florida in the north. chiropteran fauna of St. Lucia. Methods and Materials Geophysical description of the study area.—The 1766 (Lefort de Latour 1787) and a flurry of phreatic Lesser Antilles is a chain of volcanic islands that lie activity from 1839 to 1843 (Breen 1844). along the eastern edge of the Caribbean plate. This archipelago was initially formed by volcanic eruptions Saint Lucia is located in the Lesser Antilles, a more than five million years ago (Newman 1965; Lind¬ chain of islands that extends from Sombrero to the north say et al. 2002) and extends from the Greater Antilles, and to Grenada in the south. Saint Lucia is situated in nearly to the coast of South America. Islands in the the southern portion of this archipelago, 33.5 km south southern half of the Lesser Antilles were composed of Martinique and 46 km north of St. Vincent (Fig. 1). primarily of volcanic ejecta (Fig. 1). St. Lucia is 43.5 km long and 22.5 km wide. With a land area of 616 square km, it is one of the larger islands in Saint Lucia was formed approximately 2 mil¬ the Lesser Antilles. St. Lucia is characterized by very lion years ago, and its geological history has received rugged mountainous terrain, dominated by a central considerable attention (Lindsay et al. 2002). The semi¬ ridge running almost the full length of the island, slowly circular depression around the Soufriere—the Qualibou rising to Mount Gimie (958 m) in the south. From this depression—was thought by many to be an ancient central ridge, valleys extend to either coast, some of caldera (Wohletz et al. 1986). However, given that the which are broad with relatively large areas of flat land spectacular Gros and Petit Piton were the remnants of presently occupied by banana (see following section two large dacitic lava domes formed no more than 300 for scientific names of plant species discussed in text) thousand years ago, Roobol et al. (1983) and Wright plantations, including those at Cul-de-Sac and Roseau. et al. (1984) interpreted the Qualibou depression not as a caldera, but rather a large gravity slide. Another The island has a tropical marine climate charac¬ extremely violent phase of island building occurred terized by a relatively uniform dry season from January at the Soufriere Volcanic Centre 40 to 20 thousand to April and a rainy season from May to August, with years ago when a series of major eruptions produced usually sunny, warm weather from September to Oc¬ pyroclastic flows that surged southwards thus forming tober. The mean annual temperature is approximately that southern end of the island (Wright et al. 1984; 26° C at sea level. Annual rainfall varies from 1.52 to Lindsay et al. 2002). Wohletz et al. (1986) proposed 1.57 m in the north to 2.54 to 3.68 m in the mountainous that these eruptions came from the Qualibou depression interior of the south (Toussaint et al. 2009). itself, whereas others (Roobol et al. 1983; Wright et al. 1984) thought that these eruptions came from vents in Historically, tropical storms and hurricanes are the Central Highlands (Mt. Grand Magazin and Piton infrequent, with the majority passing to the north of St. Esprit). There was a massive phreatic eruption in Saint Lucia. From 1780 to 2010,16 hurricanes and 46 Pedersen et al.—Bats of Saint Lucia 3 62°0'0"W 60°0'0"W 16°0'0"N 14°0'0"N 12°0'0"N 10°0'0"N Figure 1. Map of the southern Lesser Antilles, Trinidad, and northern South America, showing the position of the island of Saint Lucia in relation to other islands and landmasses in the region. 4 Special Publications, Museum of Texas Tech University tropical storms have impacted the island. The strongest storms and hurricanes in the past. Breadfruit trees were hurricane to hit St. Lucia in recent history was Hurri¬ often seen growing in these areas along with pioneer cane Allen, a Category 4 storm (202 km/hr maximum trees such as gumtree, maho kochon, and bwa kannon. wind speed), which passed just south of the Hewanorra International Airport on 4 August 1980. St. Lucia Of lesser importance to bats are the dry scrub received catastrophic damage including: destruction woodlands that occupy the low-lying areas along the of the banana crop, extensive damage to forests, the east and west coasts and the northern part of the island. leveling of homes, and the deaths of at least 16 people. In several places, these areas are degraded through hu¬ Hurricane Dean, a Category 2 hurricane (175 km/hr man activity such as agriculture, grazing, roads, and maximum wind speed), passed between St. Lucia and settlements (Gonzalez and Zak 1996). The largest trees Martinique on 17 August 2007, after our 2007 field sea¬ in these areas are white cedar and bay leaf, but two son. Martinique sustained the most damage from this introduced trees dominate these woodlands, logwood storm, but St. Lucia lost 75% of its banana crop, which and acacia. Among the bushes and small trees common was a severe blow to the island’s economy (Caribbean in this habitat were rough-leaf velvetseed, blacktorch, Hurricane Network 2015). Natural disturbances, such and cup tree. as landslides and hurricanes, explain why relatively few of the forest areas display a classic climax structure Plant names.—The following are the common (Tomblin 1981; Daltry 2009; Graveson 2009). and scientific names of plants discussed in the text: Acacia {Acacia nilotica), Acomat Boucan {Sloanea Nearly 35% of St. Lucia is still under some form caribaea), Almond (Prunus dulcis), Balata Chien (Pou¬ of forest cover that has been described in detail (Beard ter ia pallida), banana (Musa sp.), Bay Leaf (Pimenta 1949; see also Clarke 2009; Daltry 2009; Graveson racemosa), Bitterwood (Simarouba amara), Blacktorch 2009; Morton 2009; Graveson and Smith 2013). The (Erithalis fruticosa), Blue Mahoe (Hibiscus elatus), rain forests (moist forest) covered 20% of the island Bois Cote (Tapura latifolia), Bois de Masse (Licania and occupied the higher elevations (typically above 250 ternatensis), Breadfruit (Artocarpus altilis). Butter¬ m) of the central spine of mountains extending from cup Tree (Cochlospermum vitifolium), Bwa Kannon the Castries Water Works Reserve and Piton Flore area (Cecropia schreberiana), Bwapen Mawon (Magnolia in the north across the narrow Barre de Lisle corridor dodecapetala), Candlewood (Dacryodes excelsa), to the large blocks of forest to the south exemplified Caribbean Pine (Pinus caribbea), Cocoa (Theobroma by the Edmund Forest Reserve (Isaac and Bourque cacao). Coconut (Cocos nucifera). Cup Tree (Wedelia 2001). These forests were characterized by trees calycina). Guava (Psidium guajava), Guinep (Melicoc- such as candlewood, acomat boucan, bois de masse, cus bijugatus), Gumtree (Sapium caribaeum), Lansan laurier canelle, and bwapen mawon. Plantations of (Protium attenuatum), Laurier Canelle (Phoebe elonga- non-native trees, such as blue mahoe and Caribbean ta). Limes (Citrus latifolia). Logwood (Haematoxylum pine, have been placed around the boundaries of the campechianum), Maho Kochon (Sterculia caribaea). forest reserves. Mango (Mangifera indica). Mountain Palmetto (Eu¬ terpe globosa). Nutmeg (Myristica fragrans), Pale¬ Further down the slopes was the lower montane tuvier (Tovomita plumieri), pepper plant (Piper sp.), rain forest, which was well developed on St. Lucia, with Rough-leaf Velvetseed (Guettarda sacabra), Soursop trees reaching 30 m in height in some areas. These for¬ (Annona muricata), Swizzlestick Tree (Quararibea ests were found in the lower reaches of Barre de Lisle turbinata). Tree Ferns (Cyathea arborea), and White Ridge, Piton Flore, Dennery Water Works Reserve, Au Cedar (Tabebuia heterophylla or T. pallida). Leon Peak, and Raillon Negres. Common trees of this forest type included bois de masse, balata chien, candle- Collection of specimens.—We explored and wood, bois cote, mountain palmetto, and paletuvier. collected specimens from two caves in 2007, Grace This secondary rain forest was surrounded by disturbed Cave and Soufriere Cave. However, field collection habitat associated with abandoned agricultural lands, and sampling was accomplished primarily by ground forest logged for timber, and areas damaged by tropical level mist netting. We set 127 mist nets at 21 unique Pedersen et al.—Bats of Saint Lucia 5 localities (Barre de l’Isle was sampled twice), during island-wide bat diversity and the diversity observed at four different time periods: 24-26 May 1987; 14-20 netting locations that were located away from known June 2007; 28-31 July and 1-3 August 2008; and 12-16 roosts. The latter varied amongst the three neighbor¬ March 2009. These nets were set at elevations ranging ing islands (Martinique, 24; St. Lucia, 20; and St. from 1 to 550 m and were equally distributed above Vincent, 28) as did the number of bats that were netted and below 250 m in elevation. They were also equally at these particular locations (1,859, 1,520, and 1,679, distributed on the western (wet) and eastern (dry) halves respectively). However, netting locations on wet and of the island. Netting was conducted in a variety of dry (west versus east) sides of these islands (-13:11, habitats including naturally vegetated ravines or ghuts, 10:10, 14:14) and the distributions of net sites above access roads, trails, rivers, ponds, fruit plantations, and below 250 mare comparable (12:12,10:10,14:14). forest plantations and reserves, botanical gardens, and other covered flyways. At each netting site, five to eight Minitab 16 (2010) provided standard statistics mist nets (2.8, 6, 9, 12 or 18 m length by 2.8 m height) for each sample set examined and paired t-tests were were set at 20 to 100 m intervals depending on local utilized to test for differences between groups. Species landscape features, i.e., width of access roads or rivers, curves were generated in JMP 12 (2015) and R was forest edge, vegetated dry and wet ghuts, small fruit used in the multiple regression analysis (R Develop¬ plantations, and freshwater ponds. Nets were opened ment Core Team 2008). near sunset (1800-1900 hr) and closed between 2200 and 2300 hr each night, depending on weather and Museum voucher specimens.—During the 2007- bat activity. Bats caught in nets were placed in hold¬ 2009 surveys, we captured 1,624 bats, 1,248 of which ing bags until the end of the netting each night. Bats were captured/released. We deposited 376 voucher were subsequently examined for: weight (g), length specimens in the mammal collections of the Museum of of forearm (mm), reproductive condition, tooth wear, Texas Tech University. We examined 72 bats (UNSM) presence of ectoparasites, and scars. that had been collected by two of the authors (HHG and CJP) in 1987. We examined 122 additional specimens For comparisons with our 1987 and 2007-2009 that were collected before 1987 that were deposited in collections on St. Lucia, we drew additional netting data other museums: American Museum of Natural History, from similar survey efforts on both Martinique (Catze- 2; Kansas Museum of Natural History, 32; and National flis et al., personal communication) and St. Vincent Museum of Natural History, 88 (see Appendix). There (Kwiecinski et al. 2018). These data were parsed into are no fossil records of bats from this island. Systematic Accounts The species accounts that follow are arranged in (mm). All measurements of embryos are crown-rump systematic order (Simmons 2005; Baker et al. 2016; length (mm). Testes were measured for length (mm). Cirranello et al. 2016). Given that the nine genera of Distances were recorded in kilometers (km) or miles as bats recorded from St. Lucia are monotypic on the is¬ they appeared on original specimen tags. All elevations land, we refer to these taxa by genus, where pragmatic. are reported in meters (m), unless specified otherwise. These data are drawn from both the authors’ inventory Length of forearms and cranial measurements were efforts on St. Lucia and data from museum specimens. taken from museum vouchers following Hall (1946), A list of specimens and a list of additional records are except greatest length of skull included incisors, and included. Weights were determined with a digital bal¬ length of forearm was the distance from the olecranon ance and recorded in grams (g). Forearms were mea¬ process of the elbow joint to the tip of the carpals with sured with digital calipers and recorded in millimeters wing in the retracted position. 6 Special Publications, Museum of Texas Tech University Family Noctilionidae Lesser Antillean populations ofV. leporinus. A putative Noctilio leporinus mastivus (Vahl, 1797) hybrid zone between N. 1. mastivus and N. 1. leporinus Greater Fishing Bat was identified by Khan et al. (2014) in eastern Ven¬ ezuela and the Guyana Shield, the regions from which Specimens examined(6).—Anse La Raye: Anse southern Lesser Antillean populations presumably La Raye, 1 m [13°56'30.3"N, 61°02'28.2"W], 4 (TTU originated (Genoways et al. 2010; Khan et al. 2014). 110080-83). Micoud: Canelles River, 1.5 km S Anse Additional research will be required to determine the Ger, 10 m [13°46'59.6"N, 60°54'53.1"W], 1 (TTU extent of hybridization in mainland populations of N. 110084). No Specific Quarter: no specific locality, 1 leporinus and whether or not this hybrid zone has in¬ (NMNH 110922). fluenced southern Lesser Antillean populations. With respect to Caribbean N. leporinus, we recommend the Additional records.—Soufriere: Edmund For¬ usage of advanced genomic techniques to examine the est Reserve (Clarke 2009). No Specific Quarter: no evolutionary history of Greater Antillean and Lesser specific locality (Allen 1911). Antillean populations and to more precisely identify zones of secondary contact between N. 1. mastivus and This species represents a recent radiation within N. 1. leporinus. Until such data become available, we the genus Noctilio, having diverged from its sister spe¬ provisionally retain N. 1. mastivus for the St. Lucia cies N. albiventris within the last million years (Pavan population. et al. 2013; Khan et al. 2014). Owing to the species’ recent evolutionary history, studies focused on the Table 1 presents length of forearm and seven intraspecific morphological and genetic variation of cranial measurements for four males and two female N. leporinus have resulted in conflicting taxonomic Noctilio from St. Lucia. As is typical for this spe¬ assessments. Davis (1973) identified three subspecies cies, males averaged larger than females for all seven within N. leporinus—N. 1. leporinus (distributed in measurements. The average forearm and five of seven South America east of the Andes Mountains, through¬ cranial measurements were not significantly different out Amazonia and the Guyana Shield); N. 1. mastivus between the sexes. The zygomatic breadth and length (distributed throughout Central America, the Carib¬ of maxillary toothrow were significantly (P < 0.05) bean, and in South America throughout Venezuela and larger in males than females. In comparison with along the western versant of the Andes Mountains into measurements from other West Indian females (Davis northern Ecuador), and N. 1. rufescens (distributed in 1973), measurements for females from St. Lucia were southern South America throughout Bolivia, Paraguay, within ranges reported for the West Indies except northern Argentina and southern Brazil). However, ge¬ breadth across the upper molars was smaller in the St. netic data presented in Pavan et al. (2013) and Khan et Lucia sample. al. (2014) reveal two main lineages within N. leporinus, one corresponding to N. 1. mastivus and the other to This species is widespread in the Antilles, being N. 1. leporinus. To date, all specimens collected from found in non-arid lowland and coastal regions and in within the range of N. 1. rufescens are genetically in¬ major river basins (Hood and Jones 1984). H. S. Branch distinguishable from N. 1. leporinus (Pavan et al. 2013; collected the oldest known specimen of Noctilio from Kahn et al. 2014). St. Lucia in 1901 (NMNH 110922) from an unspeci¬ fied locality. We collected Noctilio on St. Lucia at two Phylogeographic data from N. leporinus indicates localities, foraging over a coastal inlet in Anse La Raye dual invasions into the Caribbean, with the Greater An¬ on the west coast and the Canelles River on the east tilles being colonized by Central American populations coast (Fig. 2). and the Lesser Antilles colonized by northern South American populations (Lewis-Oritt et al. 2001a, 2001b; At Anse La Raye, we placed three mist nets over Genoways et al. 2010; Pavan et al. 2013; Kahn et al. the Petite Riviere de l’Anse La Raye, which was ap¬ 2014). There was no clear genetic consensus regard¬ proximately 4 m wide, at the point it crossed under the ing the appropriate subspecific taxonomy for southern coastal highway at the north edge of town (Fig. 3). Two