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Schefflera Morototoni Trees in Puerto Rico1

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Schefflera Morototoni Trees in Puerto Rico1 BIOTROPICA 37(1): 81–87 2005 Crop Size and Fruit Neighborhood Effects on Bird Visitation to Fruiting Schefflera morototoni Trees in Puerto Rico1 James F. Saracco2, Jaime A. Collazo, Martha J. Groom3, and Tomas´ A. Carlo4 North Carolina Cooperative Fish and Wildlife Research Unit, Biological Resources Division, U. S. Geological Survey, Department of Zoology, North Carolina State University, Raleigh, North Carolina 27695-7617, U.S.A. ABSTRACT Studies of zoochorous seed dispersal systems often consider crop size, yet seldom consider the kinds and amounts of fruits surrounding parent plants (the fruit neighborhood) when attempting to explain among-plant variation in fruit removal. We studied avian frugivory at 24 Schefflera morototoni trees from February to May 1998 in central Puerto Rico. The number of fruits removed by avian seed dispersers per visit was similar among focal trees (typically 2–4). In contrast, visitation rate was highly variable (range: 0–71 visits per 4 h). We used multiple regression analyses to evaluate the relative roles of crop size (focal tree ripe fruit abundance) and fruit neighborhood variables (measured within 30 m of focal trees) in affecting visitation to focal trees by avian frugivores. Visitation rate was positively related to crop size (although this variable was only significant in one of four regression models considered) and negatively related to the presence or abundance of conspecific fruits, suggesting that trees competed intraspecifically for dispersers. Relationships between visitation and heterospecific fruits were mixed—some kinds of fruits appeared to enhance visitation to focal trees, while others seemed to reduce visitation. In most regression models, neighborhood variables had larger effects on visitation than focal tree fruit crop size. Our results highlight the important effects of local fruiting environments on the ability of individual plants to attract seed dispersers. RESUMEN Los estudios de sistemas dispersadores de semillas zoocoras por lo general consideran el tamano˜ de la cosecha, sin embargo, raras veces consideran los tipos y cantidades de frutos en los alrededores de las plantas madres (las frutas del vecindario) cuando intentan explicar entre plantas, la variacion´ de extraccion´ de frutos. Nosotros estudiamos la frugivoria de aves en 24 arboles´ de Schefflera morototoni entre Febrero y Marzo de 1998 en la parte central de Puerto Rico. El numero´ de frutos removidos en cada visita por las aves dispersadoras de semillas fue similar entre los arboles´ focales (t´ıpicamente de 2 a 4). En contraste, la proporcion´ de visitas fue´ muy variable (intervalo = 0–71 visitas cada 4 horas). Nosotros utilizamos analisis´ de regresion´ multiple´ para evaluar el rol relativo del tamano˜ de la cosecha (arboles´ focales con abundancia de frutos maduros) y variables en frutos del vecindario (medidos dentro de los 30 m de los arboles´ focales) en el efecto de visitas por aves frug´ıvoras a los arboles´ focales. La proporcion´ de visitas estuvo positivamente relacionado con el tamano˜ de la cosecha (aunque esta variable fue´ solamente significativa en uno de los cuatro modelos considerados) y negativamente relacionado a la presencia y a la abundancia de frutos coespec´ıficos, sugiriendo que los arboles´ compitieron intraespec´ıficamente por los dispersores. Las relaciones entre visitas y frutos heteroespec´ıficos fue mixta -algunos tipos de frutos parecen promover las visitas a los arboles´ focales, mientras que otros parecen reducir las visitas. En la mayor´ıa de los modelos de regresion,´ las variables de “vecindario” tuvieron un mayor efecto sobre las visitas que en el tamano˜ de la cosecha de frutos del arbol´ focal. Nuestros hallazgos enfatizan la importancia que pueden tener los efectos de los ambientes locales de frutos en la habilidad de plantas individuales en atraer dispersadores de semillas. Key words: frugivory; plant–animal interactions; seed dispersal; Spindalis portoricensis; shade coffee plantation; tropical forest. MANY PLANT SPECIES PRODUCE FLESHY OR ARILLATE FRUITS that are dis- that consistently explains large proportions of among-plant variation in persed by animals (Howe & Smallwood 1982). Identifying factors that fruit removal is fruit quantity (i.e., crop size; Davidar & Morton 1986, influence plant choice by frugivores is a critical step in linking frugi- Denslow 1987, French et al. 1992, Willson & Whelan 1993, Alcantara´ vore behavior and plant demography, and thus in understanding how et al. 1997, Jordano & Schupp 2000). Nevertheless, frugivore responses animals affect the distribution, abundance, and evolution of plant mu- to crop size can be highly variable over space and time (e.g., Sallabanks & tualists (Schupp 1993). Most studies attempting to make this link have Courtney 1993, Ortiz-Pulido & Rico-Gray 2000). Much of the spatial focused on characteristics of parent plants (i.e., fruit quality or quantity). and temporal variation in visitation and fruit removal rates among plants In controlled aviary experiments, frugivorous birds often discriminate could be due to factors external to parent plants, yet these variables are among fruits that vary in aspects of fruit quality, such as color, nutrients, seldom considered. or accessibility (e.g., Moermond & Denslow 1983, Johnson et al. 1985, Among the most likely external variables to influence avian foraging Stiles 1993, Siitari et al. 1999, Schaefer et al. 2003). Such selectivity has are the kinds and amounts of fruits available to frugivores within some proven difficult to show in the field, where the only plant characteristic area surrounding parent plants (the “fruit neighborhood”; Herrera 1986). A number of studies have shown fruit removal rates at individual plants to 1 Received 19 February 2004; revision accepted 9 July 2004. be negatively related to the presence or abundance of nearby conspecific 2 Current addresses: The Institute for Bird Populations, P.O. Box 1346, Pt. Reyes (Moore & Willson 1982, Manasse & Howe 1983, Denslow 1987; but see Station, CA 94956 U.S.A., and Biology Program, University of Alaska Southeast, 11120 Glacier Highway, Juneau, AK 99801, U.S.A.; e-mail: [email protected] French et al. 1992) or heterospecific (Herrera 1984) fruits. Nevertheless, 3 Current addresses: Interdisciplinary Arts and Sciences, University of Washington, a plant’s ability to attract seed dispersers could also be enhanced by the Bothell, WA 98021-4900, U.S.A., and Department of Biology, University of presence of nearby fruiting trees if these neighbors attract frugivores to Washington, Seattle, WA 98195-1800, U.S.A. 4 Current address: Department of Ecology and Evolutionary Biology, University of areas that might not otherwise be visited (Sargent 1990, Garc´ıa et al. Colorado, Boulder, CO 80309-0334, U.S.A. 2001, van Ommeren & Whitham 2002). Such facilitative interactions 81 82 Saracco, Collazo, Groom, and Carlo between plants could be particularly prevalent on small spatial scales or S. morototoni fruits are 2-seeded drupes (4–6 mm long, 6–10 mm where fruit is especially scarce or patchily distributed (Rathcke 1983). broad) borne on 20–60 cm compound panicles (Liogier 1995). The In this paper, we describe interactions between avian frugivores and pulp is relatively rich in lipids and proteins (Snow 1971, Herrera 1981). fruiting plants during a single fruiting season in central Puerto Rico. For Crop sizes observed during the study averaged about 68,000 fruits; the the most commonly consumed plant species, Schefflera morototoni,weuse largest tree contained an estimated 1,000,000 fruits (estimation method multiple regression models to assess variation in bird visitation rates to in- described below in “Crop size and neighborhood measurements”). It was dividual fruiting trees as a function of crop size (ripe fruit abundance) and the most abundant fruit in the study area (Carlo et al. 2003). Fruits the presence or abundance of nearby ripe conspecific and heterospecific ripen in almost every month in Puerto Rico. During 1998, ripe fruits fruits. We expected visitation rates to be most strongly influenced by in- were primarily available between February and June on the study area, traspecific interactions among plants (Howe & Estabrook 1977); that is, peaking in abundance between March and April. by crop size and conspecific neighborhood fruits. Furthermore, assuming that seed-dispersing frugivores were a limiting resource to fruiting trees, PLOT-LEVEL FORAGING OBSERVATIONS.—To identify plant species likely we expected these intraspecific interactions to be competitive. Because to influence bird visitation to S. morototoni trees, we conducted foraging heterospecific fruits were less abundant and less commonly consumed by observations within a centrally located 8.28 ha grid between 2 February avian frugivores, we expected them to be of lesser importance in explain- and 18 May 1998 (data are a subset of those reported in Carlo et al. ing focal tree visitation rates. Furthermore, it has been suggested that 2003). On 6–7 days each month, one observer traversed the grid for many bird-consumed fruits may be complementary resources (Whelan approximately 5 h, beginning about 1 h after sunrise. On each observa- et al. 1998). Thus, we expected interactions between focal trees and at tion day, the observer commenced searching for foraging birds from a least some heterospecific fruit taxa would be facilitative in nature. random starting point. Movement from the starting point was guided by visual and auditory cues of seven fruit-eating bird species:
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