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Caribbean Naturalist No. 35 2016 Conservation Value of Remnant Habitat for Neotropical Bats on Islands Armando Rodríguez-Durán and Waldemar Feliciano-Robles The Caribbean Naturalist . . . ♦ A peer-reviewed and edited interdisciplinary natural history science journal with a re- gional focus on the Caribbean ( ISSN 2326-7119 [online]). ♦ Featuring research articles, notes, and research summaries on terrestrial, fresh-water, and marine organisms, and their habitats. The journal's versatility also extends to publishing symposium proceedings or other collections of related papers as special issues. ♦ Focusing on field ecology, biology, behavior, biogeography, taxonomy, evolution, anatomy, physiology, geology, and related fields. Manuscripts on genetics, molecular biology, anthropology, etc., are welcome, especially if they provide natural history in- sights that are of interest to field scientists. ♦ Offers authors the option of publishing large maps, data tables, audio and video clips, and even powerpoint presentations as online supplemental files.

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Cover Photograph: Stenoderma rufum (Red Fruit Bat). Photograph © Waldemar Feliciano. CARIBBEAN NATURALIST Board of Editors James D. Ackerman, Department of Biology, University of Puerto Rico at Río Piedras, USA Alfonso Aguilar-Perera, Department of Marine Biology, Universidad Autónoma de Yucatán, Mexico Wayne J. Arendt, International Institute of Tropical Forestry, Luquillo, Puerto Rico, USA Rüdiger Bieler, Field Museum of Natural History, Chicago, IL, USA Christopher P. Bloch, Department of Biological Sciences, Bridgewater State University, Bridgewater, MA, USA William R. Buck, Institute of Systematic Botany, New York Botanical Garden, Bronx, NY, USA Leo Douglas, Department of Geography/Geology, University of the West Indies, Mona, Jamaica Robert Erdman, Department of Biological Sciences, Florida Gulf Coast University, Fort Myers, FL, USA Keith Goldfarb, GoldRush Science Services, Steuben, ME, USA ... Editor-in-Chief Grizelle González, International Institute of Tropical Forestry, San Juan, Puerto Rico, USA Gary R. Graves, Department of Vertebrate Zoology, Smithsonian Institution, Washington, DC, USA S. Blair Hedges, Department of Biology, Pennsylvania State University, University Park, PA, USA Julia A. Horrocks, Dept. of Biological and Chemical Sciences, Univ. of the West Indies, Cave Hill Campus, Barbados Scott Jones, Smithsonian Institution, Caribbean Coral Reef Ecosystems, Carrie Bow Cay, Belize Heather Judkins, Department of Biological Sciences, University of South Florida, St. Petersburg, FL, USA Craig A. Layman, Department of Applied Ecology, North Carolina State University, Raleigh, NC, USA John Leavengood, Department of Entomology, University of Kentucky, Lexington, KY, USA Joerg-Henner Lotze, Eagle Hill Institute, Steuben, ME, USA ... Publisher Antonio A. Mignucci-Giannoni, Manatee Conservation Center, Inter American University, Bayamón, Puerto Rico, USA Gregg Moore, Department of Biological Sciences, Jackson Estuarine Laboratory, University of New Hamp- shire, Durham, NH, USA Dawn Phillip, Department of Life Sciences, University of The West Indies, St. Augustine, Trinidad and Tobago... Managing Editor James Pitts, Department of Biology, Utah State University, Logan, UT, USA Robert Powell, Department of Biological Sciences, Avila University, Kansas City, MO, USA Chris Rimmer, Vermont Center for Ecostudies, Norwich, VT, USA Armando Rodríguez-Durán, Dean for Research, Inter American University, Bayamón, Puerto Rico, USA Noris Salazar Allen, Smithsonian Tropical Research Institute, Panama Inés Sastre de Jesus, Biology Department, University of Puerto Rico at Mayagüez, USA Amy Siuda, Collegium of Natural Sciences, Eckerd College, St. Petersburg, FL, USA J. Angel Soto-Centeno, American Museum of Natural History, Division of Mammalogy, New York, NY, USA Christopher Starr, Department of Life Sciences, University of the West Indies, St. Augustine, Trinidad and Tobago David W. Steadman, Florida Museum of Natural History, Gainesville, FL, USA Kathleen Sullivan Sealey, Department of Biology, University of Miami, Coral Gables, FL, USA Jarrod M. Thaxton, Department of Biology, University of Puerto at Mayagüez, USA Jason M. Townsend, Hamilton College, Biology Department, Clinton, NY, USA Jill Weber, Eagle Hill Institute, Steuben, ME, USA ... Production Editor Byron Wilson, Department of Life Sciences, University of the West Indies at Mona, Kingston, Jamaica Graham A. J. Worthy, Department of Biology, University of Central Florida, Orlando, FL, USA Joseph M. Wunderle, International Institute of Tropical Forestry, University of Puerto Rico at Río Píedras, USA

The Caribbean Naturalist (ISSN # 2326-7119) is published by the Eagle Hill Institute, PO Box 9, 59 Eagle Hill Road, Steuben, ME 04680-0009. Phone 207-546-2821, FAX 207-546-3042. E-mail: [email protected]. Webpage: www.eaglehill.us/cana. Copyright © 2016, all rights reserved. Periodical postage paid in Steuben, ME and additional mailing offices. Special issue proposals are welcome. On-line secure subscription ordering: rate per year for Caribbean subscribers - $15 regular, $10 students, $70 organizations; for Non-Caribbean subscribers - $20 regular, $15 students, $90 organizations. Authors: submission guidelines are available at www. eaglehill.us/cana. Co-published journals: The Northeastern Naturalist (ISSN 1092-6194 [print], ISSN 1938-5307 [online]), the Southeastern Naturalist (ISSN 1528-7092 [print], ISSN 1938-5412 [online]), and the Urban Naturalist (ISNN #2328-8965), journals with separate Boards of Editors. The Eagle Hill Institute is a tax exempt 501(c)(3) nonprofit corporation of the State of Maine (Fed- eral ID # 010379899). 2016 CARIBBEANCaribbean Naturalist NATURALIST No. 35:1–10No. 35 A. Rodríguez-Durán and W. Feliciano-Robles

Conservation Value of Remnant Habitat for Neotropical Bats on Islands

Armando Rodríguez-Durán1,* and Waldemar Feliciano-Robles1

Abstract- The archipelago of remnant habitats that results after a fragmentation event often vary in the quality of ecological characteristics that allow each fragment to sustain populations of bats. The species diversity and richness of these forest remnants are influenced also by the permeability of the matrix on which they stand. Urban areas represent an ever-increasing matrix in the tropics, interspersed with remnants of natural habitats. We conducted a sampling program at 2 locations with different levels of urban encroachment along the karst belt of northern Puerto Rico. Field work was conducted from January 2013 through August 2015. During this study, we captured a total of 11 out of the 13 species of bat documented as present in Puerto Rico. Neither weather conditions nor the phase or presence of the moon were statistically significant factors determining the number of bats captured per night. Our results support the prediction that some forest remnants within an urban matrix have conservation value for bats, and that higher landscape complementation may have a positive effect on bats’ species richness and diversity. This effect may be augmented on islands, where edge-tolerant species seem to be more common than in the mainland. The permeability of the urban matrix, combined with resident populations and a network of corridors, convey important conservation value to archipelagos of forest remnants within this urban matrix.

Introduction The fragmentation of natural ecosystems is a growing reality in an epoch that, some propose, should be named the Anthropocene (Crutzen and Stoermer 2000). Habitat fragmentation and habitat reduction represent 2 results of the human modi- fication of ecosystems that are often confounded (Ewers and Didham 2006, Fahrig 2003). Whereas fragmentation per se may not have detrimental effects on bat diversity and richness (Ethier and Fahrig 2011), it is often associated with habitat reduction (e.g. Benchimol and Peres 2015, Cosson et al. 1999), which does have detrimental effects (Gibson et al. 2013, Lewis et al. 2015). After a fragmentation event, the resulting archipelago of habitats often varies in the quality of ecological characteristics that allow it to sustain populations of bats (e.g., Bernard and Fen- ton 2007, Estrada-Villegas et al. 2010, Pina et al. 2013). The species diversity and richness of these habitat “islands” is influenced not only by their intrinsic characteristics, but also by the permeability of the matrix on which they stand. Water and agricultural fields are 2 of the matrices that have been examined more thoroughly (e.g., Cosson et al. 1999, Estrada et al. 1993, Meyer and Kalko 2008, Ripperger et al. 2014). However, accelerated urbanization, with its concomitant problems, can

1Universidad Interamericana, Departamento de Ciencias Naturales, 500 John W. Harris Road, Bayamón, PR 00957. *Corresponding author - [email protected]. Manuscript Editor: J. Angel Soto-Centeno 1 2016 Caribbean Naturalist No. 35 A. Rodríguez-Durán and W. Feliciano-Robles be increasingly seen throughout the tropics (Wright 2005), making urban areas a growing matrix interspersed with remnants of natural habitats (Ferreira et al. 2010). Although not universally applicable to every taxa, certain organisms, including bats, can persist in fragments of forests (Meyer and Kalko 2008, Turner and Corlett 1996). This has probably been the case for Hong Kong and Puerto Rico, which retained a large percentage of their biodiversity despite near total deforestation during the past centuries (Lugo et al. 2001, Turner and Corlett 1996). With an approximate surface area of 8900 km2, Puerto Rico is one of the major islands of the West Indian archipelago in the Caribbean. Extensive deforestation of the island started during the 19th century, and by 1940, forests represented just 6% of its surface. By 1985, forest cover had increased to 34% for the island as a whole, and to 49% for the northern karst belt (Birdsey and Weaver 1987). However, from 1977 to 1994, the amount of urbanized land increased over 25% (López et al. 2001). This accelerated urbanization occurred mostly along the coastal plains, leaving a number of forest fragments encroached by an urban matrix. Here, we examine 2 localities along the northern karst belt of Puerto Rico to assess the conservation value of habitat remnants within an urban matrix. Given the increasing urban development in the Neotropics (Wright 2005), this case could serve as a platform to understand the consequences of urban encroachment else- where. Based on a previous study looking at a habitat impacted by agricultural activities and moderate urban development (Rodríguez-Durán and Otero 2011), we predicted that forest remnants within an urban matrix would have a medium to high conservation value.

Field-Site Description We conducted a sampling program at 2 locations, ~50 km apart, along the karst belt of northern Puerto Rico. The main urban area of Puerto Rico, the San Juan Metropolitan Area (SJMA), is located along the eastern portion of the karst belt. The central part of the belt constitutes the largest tract of continuous forest cover on the Island (Lugo et al. 2001). We selected 1 location in an area protected by the Puerto Rico Conservation Trust, within the municipality of Ciales in the central forested region of the karst belt, which we designated as the wilderness location (WS). The second location is an urban park (230 ha) situated between the municipalities of Guaynabo and Bayamón, within the SJMA, which we labeled as the remnant location (RT). Both locations were compared to Hacienda La Esperanza (HLE), also within the northern karst belt, and where an identical sampling protocol was followed (Rodríguez-Durán and Otero 2011). As with most forested areas in Puerto Rico, all localities consist of secondary growth forest, some of which can be over 60 years old. Location WS constitutes abandoned agriculture land, whereas RT was a military base from World War II and holds several abandoned underground military bunkers. There are also potential habitat corridors and “stepping stones” that connect RT to other forested areas (Fig. 1).

2 2016 Caribbean Naturalist No. 35 A. Rodríguez-Durán and W. Feliciano-Robles

Methods We conducted field work from May 2013 through June 2015 at WS and from January 2013 through August 2015 at RT, for a total of 26 consecutive months of sampling at WS and 32 at RT. We set 72 m of mist nets, 2.5 m high, every other week during the study period in a standardized manner, in a zig-zag pattern along trails with a closed canopy. The location of mist nets changed regularly, but always fell within the same general area. We used aerial photographs from Google Earth to determine the percentage of forest cover within a radius of 5 km from the netting area. Nets were opened at sunset and remained open for 4 hours. We noted weather conditions and moon phase for each sample night. In addition to mist netting, we conducted acoustic monitoring during most of the sampling to detect aerial insectivores that could be avoiding the nets. We set an ANABAT Ultrasound System on an open space about 150 m from where the nets were located. We compared calls recorded with the ANABAT to an unpublished library of calls from the bats of Puerto Rico using ANALOOK. We used multiple regression analysis (STATA®) to assess the influence of moon phase, weather, and seasonality on the number of bats captured per night. Thor- ough knowledge about the local bat fauna (Gannon et al. 2005) allowed the species

Figure 1. Map of the urban matrix where the RT site (marked black) is located. Striped areas represent non-urbanization, including secondary forest, streams with heavily impacted riparian vegetation, and abandoned pasture. 3 2016 Caribbean Naturalist No. 35 A. Rodríguez-Durán and W. Feliciano-Robles accumulation curve to be compared to known bat assemblages rather than to a predicted estimate. We calculated the percentage of species shared among locations as c/a, where c = species common to both locations and a = all species present at both locations. This comparison was also made with the HLE location. To provide a basis for comparison with previous studies (e.g., Fenton et al. 1992, Rex et al. 2008), we calculated the Shannon-Wiener index (HS), evenness (J'), and dominance using EXCEL.

Results During this study, we captured a total of 875 bats, representing 11 out of the 13 species documented as present on Puerto Rico (Gannon et al. 2005). The rate of capture (0.04 bats/hr-m of mistnetting) and the total number of captures (565 bats) was highest at WS, which has the lowest area of urban development (<5%) within a 5-km radius of the sampling site. Our RT site showed half the rate of capture (0.02 bats/hr-m) for a total of 310 captures, with >95% of urban development within a 5-km radius of the sampling site. Table 1 shows the species and number of individuals captured with mist nets at each site. Both locations show the same number of species (9) when both mist netting and acoustic monitoring are taken into consider- ation. However, species composition was different, with Brachyphylla cavernarum (Antillean Fruit-eating Bat) and Mormoops blainvillei (Antillean Ghost-faced Bat) captured at WS but not at RT, whereas Eptesicus fuscus (Big Brown Bat) and Noc- tilio leporinus (Greater Bulldog Bat) were captured at RT but not at WS. At each location, all species of insect-eating bats that were captured with mist nets were

Table 1. List of species of bats present in Puerto Rico and known to occur in the study sites. Number of captures with mist-nets is indicated for the wilderness (WS) and remnant (RT) locations, AD stands for acoustically detected and indicates species also detected with the ANABAT.

Species WS RT Insectivores Eptesicus fuscus (Palisot de Beauvois) (Big Brown Bat) 0 26 (AD) Lasiurus minor Miller (Minor Red Bat) 0 0 Molossus molossus (Pallas) (Velvety Free-tailed Bat) 0 (AD) 2 (AD) Mormoops blainvillei Leach (Antillean Ghost-faced Bat) 10 (AD) 0 Pteronotus parnellii Gray (Parnell’s Mustached Bat) 5 (AD) 9 Pteronotus quadridens (Gundlach) (Sooty Mustached Bat) 24 (AD) 1 (AD) Tadarida brasiliensis (I. Geoffroy) (Mexican Free-tailed Bat) 0 0 Piscivores/Insectivores Noctilio leporinus (L.) (Greater Bulldog Bat) 0 19 (AD) Frugivores Artibeus jamaicensis Leach (Jamaican Fruit Bat) 312 32 Brachyphylla cavernarum Gray (Antillean Fruit-eating Bat) 33 0 Stenoderma rufum Desmarest (Red Fruit Bat) 40 13 Nectivores/Frugivores Erophylla bombifrons Miller (Brown Flower Bat) 90 26 Monophyllus redmani Leach (Leach’s Single Leaf Bat) 51 182 Total 565 310

4 2016 Caribbean Naturalist No. 35 A. Rodríguez-Durán and W. Feliciano-Robles also detected with the ANABAT (Table 1). Molossus molossus (Velvety Free-tailed Bat) was captured with mist nets at RT, but only detected with the ANABAT at WS. Abandoned bunkers at RT revealed occasional bats roosting during the day, but were commonly used as night roosts by large numbers of bats. Based on the species captured with mist nets and detected with ANABAT, 63% of the species are shared by WS and RT. Each location shared 75% of the species with HLE, a third location with moderate impact from agriculture and urbanization that is located within the same northern karst belt (Rodríguez-Durán and Otero 2011). The Shannon-Wiener index of diversity based on bats captured with mist nets was similar at both locations (HS = 1.42 at RT vs 1.44 at WS); the same holds true for evenness (0.65 for WS, 0.64 for RT) and dominance (0.35 for WS, 0.38 for RT). Species were accumulated at a higher rate in our wilderness site (WS) than in the remnant forest (RT) (Fig. 2).

Figure 2. Species-accumulation curves of bats of WS (solid line) and RT (dotted line) sites as a function of total number of individuals captured (A) and months of capture (B). 5 2016 Caribbean Naturalist No. 35 A. Rodríguez-Durán and W. Feliciano-Robles Seasonality was a statistically significant variable influencing bat captures (t = 2.24, P = 0.028). Noctilio leporinus was typically captured during the first 5 months of each year, after which it virtually disappeared from the RT site. At this same location, Artibeus jamaicensis (Jamaican Fruit Bat) was captured in greater numbers during the second half of the year. At WS, we captured Monophylus redmani (Leach’s Single Leaf Bat) and Erophylla bombifrons (Brown Flower Bat) in greater numbers during the first 3 months of each year. Neither weather conditions (t = -0.59, P =0.56) nor the phase or presence of the moon (t = 0.43, P = 0.67) were statistically significant factors determining the number of bats captured per night.

Discussion Our results support the prediction that some forest remnants within an urban matrix have conservation value for bats. It also supports the suggestion that higher landscape complementation (sensu Ethier and Fahrig 2011) may have positive effects on bats’ species richness and diversity. This correlation may partly explain the high diversity of bats reported by Ferreira et al. (2010) in urban parks in Brazil. However, the effect may be augmented on islands, where edge-tolerant species are perhaps more common than on the continent (Rodríguez-Durán and Kunz 2001). Bat faunas on islands tend to have better dispersal capacity than a random sample of mainland species, and are likely to be better suited to deal with fragmented ecosystems (Meyer and Kalko 2008). Species in the Caribbean islands are subject to regular disturbances caused by hurricanes (Gannon and Willig 2009, Jones et al. 2001, Pedersen et al. 2009, Rodríguez-Durán and Vázquez 2001), which will often leave a landscape of patchy resources amongst a matrix of devastated forest. The ability of bats to use these resources may ultimately determine their capacity to survive on the islands (Rodríguez-Durán 2009). Out of the 13 species of bats on Puerto Rico, we captured a total of 11 during this study. Our results support previous studies that found no lunar phobia for bats in Puerto Rico (Gannon and Willig 1997, Rodríguez-Durán and Vázquez 2001). Although the rate of capture and accumulation of species was higher at WS (Fig. 2), each of our 2 study sites show the same number of species (9, or 69% of the species present on the island) when the results from mist-netting and acoustic monitoring are combined. These findings contrast with the results obtained by Rodríguez- Durán and Otero (2011), who captured 12 (92%) of the 13 species present on the island at HLE, located between our 2 study sites within the northern karst region. The site examined by Rodríguez-Durán and Otero (2011) lies within a combined matrix of agricultural and urban landscapes, and is better connected to wilderness areas than our RT site. The relation between landscape variables and species oc- currence remain poorly known (Chambers et al. 2016). However, we propose that habitat complementation (Ethier and Fahrig 2011) may explain the differences observed among these 3 sites. The fact that Molossus molossus was captured in a mist net at RT suggests that this species roosts within this area. Molossus molossus is an insect-eating bat that typically forages high in open spaces (Gannon et al. 2005), which makes 6 2016 Caribbean Naturalist No. 35 A. Rodríguez-Durán and W. Feliciano-Robles it an unlikely candidate to be captured with mist nets. One plausible explanation for our ability to capture this species with our nets is that it uses the abandoned bunkers at RT as day roosts. This possibility is reasonable given that this bat is more commonly found in disturbed areas (Rodríguez-Durán and Feliciano-Robles 2015), where it typically roosts in anthropogenic structures (Gannon et al. 2005, Rodríguez-Durán and Christenson 2012). At WS, we only detected this species acoustically. This observation, combined with the lack of acoustic detection of the species most commonly captured, 2 phytophagous phyllostomids (Table 1), stresses the importance of using various sampling methods when assessing presence of species (Rodríguez-Durán and Feliciano-Robles 2015). Eptesicus fuscus was one of the bats recorded at RT but not at WS. This species can be found roosting in caves, which are abundant in the area where our WS sampling site was located. However, it also roosts in culverts (Gannon et al. 2005), which are ubiquitous within the SJMA where our RT sampling site was located. Moreover, E. fuscus commonly forages in forest edges (Gannon et al. 2005). Thus, the abundance of E. fuscus within our RT site could be the result of landscape complementation provided by the urban matrix combined with forest remnants and corridors (Fig. 1). Like E. fuscus, the fish-eating batN. leporinus was detected at RT but not at WS, and only seasonally. The location of the SJMA at the edge of a major estuarine area, combined with the existence of a network of potential corridors between the coast and RT (Fig. 1), most likely explains the presence of this bat at RT. Our WS site was not located near any major body of water, which may explain why N. leporinus was not detected there (Gannon et al. 2005, Rodríguez-Durán and Christenson 2012). It is not clear why this bat uses RT on a seasonal basis. The other species showing seasonal patterns are phytophagous phyllostomids, and their seasonality was most likely associated to the phenology of the forest. The 2 species recorded from WS but not from RT, B. cavernarum and M. blainvillei, are understory foragers that roost in caves (Gannon et al. 2005). Forest-in- terior species are known to be replaced by more generalist species in fragmented forests (Aguirre et al. 2003). Our results suggest that the urban matrix may be impermeable to these 2 species, since both are common and relatively abundant throughout the forested karst region (Gannon et al. 2005, Rodríguez-Durán and Christenson 2012). Both of these species were reported from the HLE location (Rodíguez-Durán and Otero 2011), possibly as a result of the presence of continuous forest in the vicinity of HLE. Estrada and Coates-Estrada (2002) suggested that bat populations benefit from the preservation of clusters of forest fragments in the vicinity of continuous forest. Stenoderma rufum is a common species, although it is not abundant; it is captured at many locations throughout Puerto Rico (Rodríguez-Durán and Christenson 2012), but never captured in large numbers. This bat does not commute long distances to foraging areas, with average home ranges of just 2.5 ha, and it roosts solitarily or in small numbers in trees (Gannon et al. 2005). Therefore, the bats captured at RT probably represent a resident population. This is not likely to be the case with Monophylus redmani, the species most frequently captured at RT. This

7 2016 Caribbean Naturalist No. 35 A. Rodríguez-Durán and W. Feliciano-Robles fruit/nectar-feeding bat roosts in large numbers in hot-caves (Rodríguez-Durán 2009), which are not present within the RT location. This bat most likely commuted from a cave located about 6 km east of RT (Gannon et al. 2005), on a forest fragment separated from RT by an urban matrix that includes expressways, and used the bunkers as night roosts while foraging. Our results show a complex dynamic of bats’ utilization of forest remnants in an urban matrix. Habitat complementation appears to play an important role in maintaining a relatively high richness and diversity of species within our RT site. The permeability of the urban matrix to species that commute long distances at high elevations (Rodríguez-Durán 2009), or use backyards and small parks in foraging trap lines, combined with resident populations and a network of corridors, convey important conservation value to archipelagos of forest remnants within this urban matrix.

Acknowledgments Luis Santiago kindly assisted with the statistical analysis. Jean M. Sandoval and Yaniré Martínez assisted with figures and provided logistic support with many of the volun- teers that participated of field activities. Two anonymous reviewers greatly improved the manuscript. This research was sponsored by the National Science Foundation (NSF) under Informal Science Education Proposal 1223882, Para la Naturaleza, and the Conservation Trust of Puerto Rico.

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