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A Brief Description of the Diet and Feeding Behavior of the Jamaican Fruit Bat (Artibeus jamaicensis) in Westmoreland Parish, Jamaica
François Fabianek

Caribbean Naturalist, No. 21 (2014): 1–6

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Caribbean Naturalist 1 F. Fabianek 22001144 CARIBBEAN NATURALIST No. 2N1o:1. –261 A Brief Description of the Diet and Feeding Behavior of the Jamaican Fruit Bat (Artibeus jamaicensis) in Westmoreland Parish, Jamaica François Fabianek1,2,* Abstract - I evaluated the diet of two colonies of Artibeus jamaicensis (Jamaican Fruit Bat) from December 2013 to January 2014, along the western seashore of Westmoreland Parish, Jamaica. Fruits of Terminalia catappa (Tropical Almond) and Adonidia merrillii (Christmas Palm) formed the bulk of the diet, followed by fruits of Manilkara zapota (Sapodilla), Melicoccus bijugatus (Guinep), and Coccoloba uvifera (Seaside Grape), in that order. I also recorded echolocation calls of Jamaican Fruit Bats consuming ripe Artocarpus altitis (Breadfruit) during eight nocturnal surveys around feeding sites. This is the first record of Jamaican Fruit Bat consuming Christmas Palm fruit and the first mention of Breadfruit consumption by a frugivorous bat in the Caribbean islands. Introduction Bats provide many services to ecosystems worldwide (Kunz et al. 2011). Among these, effective seed dispersion by frugivorous bats helps maintain diversity in tropical forests (Fleming and Heithaus 1981, Fleming and Racey 2010). Artibeus jamaicensis Leach (Jamaican Fruit Bat) is one of the most common and widely distributed species in the Neotropics. It is considered an effective seed disperser due to the large quantities of fruits that it ingests during the night, and its broad foraging range (Lobova et al. 2009). This frugivorous bat usually shows a preference for species of Ficus (figs), but becomes more generalist during periods of low fruit availability (Ortega and Castro-Arellano 2001). Up to 190 fruit species from 43 families have been reported to be consumed by this frugivorous bat species in the Neotropics (Lobova et al. 2009). The Jamaican Fruit Bat colonized the island of Jamaica in the late Pleistocene (Fleming et al. 2010), where it has since widely distributed and has probably benefited from the agricultural planting of exotic fruit trees (Genoways et al. 2005). Nevertheless, further research is needed to fully understand the ecological importance of this widespread bat species in the Caribbean and across the Neotropics. Only one mention of this bat’s diet is available from the literature for the island of Jamaica (Genoways et al. 2005). This study describes the diet and feeding behavior of the Jamaican Fruit Bat over a two-month period in Westmoreland Parish, Jamaica. 1Centre d’Étude de la Forêt et Faculté de Géographie, Foresterie et Géomatique, Université Laval, 2405, rue de la Terrasse, Québec, QC, G1V 0A6, Canada. 2Groupe Chiroptères du Québec, 734, rue Deligny, Québec, QC, G1R 3A3, Canada. *Correspondence contact - Manuscript Editor: Armando Rodríguez-Durán Caribbean Naturalist F. Fabianek 2014 No. 21 2 Methods and Materials I conducted this study in two small limestone caves that were located 3 km apart, along the western seashore cliffs of Westmoreland Parish, Jamaica (18°15'N, 78°21'W). Each cave held up to 50 Jamaican Fruit Bats, including males and females. I placed a total of 10 fruit traps to collect the various fruits and fruit pulps that were left under feeding sites of small groups of individuals. Each of these seed traps was constructed of a 1 m x 1 m plastic sheet, which was placed 30 cm above the ground surface to prevent seed predation by other mammals. I collected fruits once a week, for a period of eight weeks from December 2013 (end of the wet season) to January 2014 (beginning of the dry season). I considered each collection as a sample (n = 8) and did not observe signs of seed removal by predation from any trap. I counted and identified the consumed fruits to species and expressed the results as mean percentage volume and mean percentage of occurrence. I determined visually the percentage volume of each species of fruit within each sample. To estimate percentage occurrence, I calculated the proportion of each fruit species within each sample. I then averaged the results for the total number of samples (Table 1). I noted if the fruits that had been consumed were ripe (R) or unripe (U), and whether the entire fruit (E) was consumed or only the pulp (P). I recorded bat echolocation calls during eight nocturnal surveys at the cave entrances and around feeding sites to confirm that the individuals that were observed during feeding were Jamaican Fruit Bats. I used an external SMX-UT microphone (Wildlife Acoustics Inc., Maynard, MA, USA) mounted on a 2-m pole and connected to an Echo Meter EM3+ bat detector (Wildlife Acoustics Inc., Maynard, MA, USA) to record Jamaican Fruit Bat echolocation calls. I compared echolocation call parameters (e.g., duration, slope, minimum and maximum frequency, number of harmonics, and frequency of the maximum amplitude) from unknown bat-call species recorded during surveys with those from a personal bat-call database, including validated echolocation calls from Jamaican Fruit Bat. I visualized and identified all recorded bat echolocation calls using Batsound software (Pettersson Elektronik AB, Uppsala, Sweden; Fig. 1). Table 1. Dietary composition of Artibeus jamaicensis (Jamaican Fruit Bat) from December to January, along the western seashore of Westmoreland Parish, Jamaica (18°15'N, 78°21'W). Sample means ± SE (n = 8) are provided for each species of fruit collected in fruit traps. Ripeness: R = ripe seed, U = unripe seed. Part eaten: P = only the pulp is consumed; E = entire seed is consumed. Part Scientific name Common name Volume (%) Occurrence (%) n Ripeness eaten Terminalia catappa Tropical Almond 63.3 ± 2.1 42.3 ± 2.3 149 R P Adonidia merrillii Christmas Palm 29.2 ± 1.1 44.5 ± 2.3 160 R P Manilkara zapota Sapodilla 4.9 ± 1.9 8.0 ± 3.1 28 U E Melicoccus bijugatus Guinep 1.8 ± 0.9 3.1 ± 1.6 11 U P Coccoloba uvifera Seaside Grape 0.8 ± 0.5 2.1 ± 1.2 7 R P Caribbean Naturalist 3 F. Fabianek 2014 No. 21 Results I found six species of fruits that were consumed by the Jamaican Fruit Bat during December and January. The fruits of Terminalia catappa L. (Tropical Almond; Combretaceae; 63% volume and 42% occurrence, n = 149) and Adonidia merrillii Becc. [= Veitchia merrillii] (Christmas or Manila Palm; Arecaceae; 29% volume and 45% occurrence, n = 160) were mainly consumed, followed by Manilkara zapota (L.) P. Royen (Sapodilla; Sapotaceae; 5% volume and 8% occurrence, n = 28), Melicoccus bijugatus Jacq. (Spanish Lime or Guinep; Sapindaceae; 2% volume and 3% occurrence, n = 11), and Coccoloba uvifera (L.) Jacq. (Seaside Grape; 1% volume and 2% occurrence, n = 7) fruits (Table 1). Fruits of Tropical Almond, Christmas Palm, and Seaside Grape were ripe when consumed, while Sapodilla and Guinep were still immature. Only the pulp was removed from the aforementioned fruits, except for those of Sapodilla, which were almost completely eaten. Guinep and Seaside grape are native species of the Caribbean islands (Nellis 1994), and the other four species of fruits that were consumed by the Jamaican Fruit Bats have been introduced on the island of Jamaica. During nocturnal surveys, I recorded (Fig. 1) up to 10 Jamaican Fruit Bats flying in circles over the canopy to consume ripe Artocarpus altitis (Parkinson) Fosberg (Breadfruit; Moraceae). I observed the same pattern of intense feeding activity that was described by Lorenzo Raíces et al. (2008) on Artocarpus heterophyllus Lam. (Jackfruit) by Phyllostomus hastatus (Pallas) (Greater Spear-nosed Bat). Individual Figure 1. Spectrograms (Fast Fourier Transform size: 1024, Hanning window) on the left and power spectrum (n = 40 echolocation calls) on the right, from Artibeus jamaicensis (Jamaican Fruit Bat). The spectrograms show two echolocation calls that were recorded from individual bats flying in circles over ripe Artocarpus altitis (Breadfruit) during eight nocturnal surveys along the western seashore of Westmoreland Parish, Jamaica (18°15'N, 78°21'W). Caribbean Naturalist F. Fabianek 2014 No. 21 4 bats landed one by one on the brown-colored epicarp for up to 20 s to consume the pasty yellow-colored mesocarp (Fig. 2). Occasionally, two individuals landed on a given fruit at the same time. No more than four nights were required for about 10 individuals to consume an entire ripe Breadfruit. Figure 2. Ripe Artocarpus altitis (Breadfruit) after the second night of consumption by Artibeus jamaicensis (Jamaican Fruit Bat) along the western seashore of Westmoreland Parish, Jamaica (18°15'N, 78°21'W). Photograph © François Fabianek. Caribbean Naturalist 5 F. Fabianek 2014 No. 21 Discussion The composition of the Jamaican Fruit Bat diet likely reflects the availability of fruits during a given period of the year and in a given area (Toribio-Hernández 2013). Abundant ripe Tropical Almonds and Christmas Palm fruits were frequently observed along the western seashore of Westmoreland Parish during this study. Seaside Grape was also frequently observed, but it remained unripe at this period of the year, which may explain the low proportion of this species in the bat’s diet. Tropical Almond is also reported as a main component of the Jamaican Fruit Bat diet on the island of Puerto Rico (Rodríguez-Durán and Vazquez 2001). Consumption of the fruits of Sapodilla, Guinep, and Seaside Grape have already been reported in the literature (Lobova et al. 2009, Ortega and Castro-Arellano 2001). However, the current study is the first to record consumption of Christmas Palm fruit and Breadfruit by the Jamaican Fruit Bat in the Caribbean islands. Until now, the consumption of Artocarpus fruits by frugivorous bats remained largely anecdotal. To date, Artibeus spp. and Greater Spear-nosed Bats have only been mentioned feeding on Jackfruit (Lorenzo Raíces et al. 2008, Lobova et al. 2009). As suggested by Lorenzo Raíces et al. (2008), the deliberate consumption of Breadfruit by frugivorous bats might be associated with a period of low fruit availability. Indeed, fruit collections that were made in December and January, which is the beginning of the dry season in Jamaica and the rest of the Neotropics, are marked by a decrease in fruit availability (van Schaik et al. 1993). I am aware that bat feeding behavior was only briefly explored during this study. However, the consumption of large, sometimes seedless, fruits by frugivorous bats might be underestimated by studies relying only upon fruits that are carried to feeding sites. I would suggest that diet analyses of tropical fruit bats should be complemented by nocturnal observations of their feeding behavior to avoid bias towards small- and medium-sized fruits that could be carried to feeding sites. Acknowledgments I thank the landowners who allowed me the access to the caves located on their properties. I thank Marie-Claude Provost for her assistance in the field. I am grateful to W.F.J. Parsons (Centre d’Étude de la Forêt) and anonymous reviewers who improved the quality of this manuscript. This research was funded by the Groupe Chiroptères du Québec. Literature Cited Fleming, T.H., and E.R. Heithaus. 1981. Frugivorous bats, seed shadows, and the structure of tropical forests. Biotropica 13:45–53. Fleming, T.H., and P.A. Racey. 2010. Island Bats: Evolution, Ecology, and Conservation. University of Chicago Press, Chicago, IL, USA. Fleming, T.H., K.L. Murray, and B. Carstens. 2010. Phylogeography and genetic structure of three evolutionary lineages of West Indian phyllostomid bats. Pp. 116–150, In T.H. Fleming, and P.A. Racey (Eds.). Island Bats: Evolution, Ecology, and Conservation. University of Chicago Press, Chicago, IL, USA. Caribbean Naturalist F. Fabianek 2014 No. 21 6 Genoways, H.H., J.W. Bickham, J.R. Baker, and J.C. Phillips. 2005. Bats of Jamaica. Special Publications of The Museum of Texas Tech University 48:154. Kunz, T.H., E. Braun de Torrez, D. Bauer, T. Lobova, and T.H. Fleming. 2011. Ecosystem services provided by bats. Annals of the New York Academy of Sciences 1223:1–38. Lobova, T.A., C.K. Geiselman, and S.A. Mori. 2009. Seed dispersal by bats in the neotropics. The New York Botanical Garden, Bronx, NY, USA. Lorenzo Raíces, D.S., F. Soares Pessôa , J. Lins Luz, T. Jordão-Nogueira, C.E.L. Esbérard, and H. de Godoy Bergallo. 2008. Feeding behaviour of the bat Phyllostomus hastatus (Pallas 1767) in Jackfruit, Artocarpus heterophyllus Lamarck (Moraceae), in Ilha Grande, Rio de Janeiro State, Brazil. Revista Brasileira de zoociências 10:265–267. Nellis, D.W. 1994. Seashore Plants of South Florida and the Caribbean: A Guide to Identification and Propagation of Xeriscape Plants. Pineapple Press, Inc., Sarasota, FL, USA. Ortega, J., and I. Castro-Arellano. 2001. Artibeus jamaicensis. Mammalian species 662:1–9. Rodríguez-Durán, A., and R. Vazquez. 2001. The bat Artibeus jamaicensis in Puerto Rico (West Indies): Seasonality of diet, activity, and effect of a hurricane. Acta Chiropterologica 3:53–61. Toribio-Hernández, E. 2013. Dinámica espacio-temporal de ensambles de murciélagos frugívoros en respuesta a la disponibilidad de frutos en selvas medianas. Chiroptera Neotropical 19:1192–1197. van Schaik, C.P., J.W. Terborgh, and S.J. Wright. 1993. The phenology of tropical forests: Adaptive significance and consequences for primary consumers. Annual Review of Ecology and Systematics 24:353–377.