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Spatial Patterns of Distribution and Abundance of Harrisia Portoricensis, an Endangered Caribbean Cactus

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Journal of Plant Ecology Advance Access published March 14, 2013 Journal of Plant Ecology Spatial patterns of distribution PAGES 1–10 doi: 10.1093/jpe/rtt014 and abundance of Harrisia available online at www.jpe.oxfordjournals.org portoricensis, an endangered Caribbean cactus Julissa Rojas-Sandoval1,2,* and Elvia J. Meléndez-Ackerman2,3 1 Department of Biology, University of Puerto Rico, Río Piedras Campus, PO Box 23360, San Juan, Puerto Rico 00931-3360, USA 2 Center for Applied Tropical Ecology and Conservation, University of Puerto Rico, PO Box 70377, San Juan, Puerto Rico 00936-8377, USA 3 Department of Environmental Sciences, College of Natural Sciences, University of Puerto Rico, PO Box 70377, San Juan, Puerto Rico 00936-8377, USA *Correspondence address. Department of Botany, National Museum of Natural History, MRC-166, Smithsonian Downloaded from Institution, PO Box 37012, Washington DC 20013–7012, USA. Tel: (001) 520 425 4518; Fax: (001) 202 633 0899; E-mail: [email protected] Abstract http://jpe.oxfordjournals.org/ Aims Important findings The spatial distribution of biotic and abiotic factors may play a dom- Abundance of H. portoricensis showed strong affinities with micro- inant role in determining the distribution and abundance of plants habitat variables related to canopy structure, soil cover and light in arid and semiarid environments. In this study, we evaluated how environment. The distribution of this cactus species throughout the spatial patterns of microhabitat variables and the degree of spatial island was consistent with the spatial variation patterns of these var- dependence of these variables influence the distribution and abun- iables. In general, landscape-level analyses suggested a predictive by guest on March 15, 2013 dance of the endangered cactus Harrisia portoricensis. value of microhabitat traits for the distribution and abundance of this endangered species. For sensitive cacti species, wherein abun- Methods dance may be influenced by similar variables, these types of analy- We used geostatistical analyses of five microhabitat variables (e.g. ses may be helpful in developing management plans and identifying vegetation cover, soil cover and light incidence) and recorded the critical habitats for conservation. abundance of H. portoricensis in 50 permanent plots established across Mona Island, Puerto Rico, by the United States Department Keywords: abundance, Caribbean cactus, geostatistics, Harrisia of Agriculture Forest Service as part of the Forest Inventory and portoricensis, Mona Island, spatial correlation, spatial distribution Analysis (USDA–FIA). We also used partial Mantel tests to evaluate the relationships between microhabitat variables and abundance of Received: 4 December 2012 Revised: 26 January 2013 Accepted: H. portoricensis, controlling for spatial autocorrelation. 10 February 2013 and abundances (Holmgren et al. 2006; Reynolds et al. 2004). INTRODUCTION With ongoing regional and global changes in primary climatic One of the major goals of ecological research is to determine variables (IPCC 2001), determining the link between biotic the factors that limit the distribution and abundance of and abiotic factors with species abundance and distribution species (Elith and Leathwick 2009; Krebs 2002; Zhang et al. is a necessary step to understand how species will respond to 2012). For any species, factors limiting its distribution include these impending changes and their adaptive capacities. Spatial a combination of biotic and abiotic factors that may operate at analyses combining plant distributions with variations in different spatial and temporal scales (Collin and Glenn 1991; abiotic and biotic factors could be helpful tools for identifying Elith and Leathwick 2009; Fortin and Dale 2005; Zhang et al. these links and determining the habitats that may be most 2012). However, in extreme environments, abiotic factors suitable for species of conservation concern (Babish 2006; are thought to have a dominant role in species distributions Fortin and Dale 2005). © The Author 2013. Published by Oxford University Press on behalf of the Institute of Botany, Chinese Academy of Sciences and the Botanical Society of China. All rights reserved. For permissions, please email: [email protected] Page 2 of 10 Journal of Plant Ecology Cacti are often a dominant component of plant communi- size of Harrisia on Mona Island using an island-wide census ties in arid and semiarid ecosystems. For species within this and a geostatistical approach, (ii) explore the spatial patterns family, low temperatures are often the single most impor- of particular microhabitat variables (i.e. canopy cover; propor- tant element limiting their latitudinal distribution (Godínez- tions of rock, soil and litter; and light incidence) and evaluate Alvarez et al. 2003; Medel-Narvaez et al. 2006). Nevertheless, how these might be associated with the abundance and dis- it is the spatial variability in a combination of factors (i.e. tem- tribution of Harrisia and (iii) determine habitat characteristics perature, topography, soil properties, rainfall and availability that would be most suitable for the establishment and growth of nurse plants) that often determines their local patterns of Harrisia. We hypothesized that the distribution and abun- of distribution and abundance (Godínez-Alvarez et al. 2003; dance of Harrisia on Mona Island would be closely associated Parker 1988). At local scales, cacti abundance may be highly with spatial patterns of microhabitat variables, principally variable, ranging from a few individuals to thousands per hec- canopy cover and light incidence. In this regard, we expected tare (de Viana et al. 1990; Esparza-Olguín et al. 2002; Medel- that areas with intermediate values of canopy cover and light Narvaez et al. 2006; Valiente-Banuet and Ezcurra 1991), and incidence would be optimal for the presence and abundance to date, most species studied exhibit clumped distributions of Harrisia. Given the protected status of Harrisia, this infor- (Fleming and Valiente-Banuet 2002). Many authors suggest mation will be valuable in identifying suitable habitats for the that local distribution and abundance patterns of cacti may conservation and management of this species. be explained by patchy distributions of environmental condi- tions that enhance germination, seedling establishment and Downloaded from plant growth (Bowers 1997; Valverde et al. 2004). However, MateriaLS AND METHODS we still lack landscape-level information about how spatial Study site and study species patterns of environmental variables may affect the final dis- This study was conducted on Mona Island, a 5517-ha raised tribution and abundance of cacti. platform of limestone rock located in the Caribbean Sea Harrisia portoricensis (hereafter, Harrisia) is a columnar cac- between Puerto Rico and the Dominican Republic (18°05′N, http://jpe.oxfordjournals.org/ tus formerly endemic to the Puerto Rican Archipelago and 67°54′W; Fig. 1). The surface of Mona Island is character- currently restricted to the small islands of Mona, Monito and ized by exposed rocks, cracks and interspersed sinkholes with Desecheo (United States Fish and Wildlife Service 1990). The accumulated soil. Water on the island is limited to organisms largest population of Harrisia occurs on Mona Island, where because the limestone rock structure ensures the rapid per- a previous study yielded a population size estimate of 59 857 colation of water and the shallow soils have low water-hold- (standard error, SE = 1058) plants (Rojas-Sandoval 2010; ing capacities (Lugo et al. 2001). The island has an annual Rojas-Sandoval and Meléndez-Ackerman 2013). The demo- mean temperature of 26.8°C and an average annual rainfall graphic profile of this population included plants in all life- of 895.79 mm. Its climate is considered semiarid because the by guest on March 15, 2013 history stages (i.e. seedlings, juveniles and adults), indicating rainfall received is significantly less than the potential evap- that at least some recruitment is occurring at this locality. otranspiration level throughout the year (Rojas-Sandoval Recent data on the establishment and growth of early life- 2010; Rojas-Sandoval and Meléndez-Ackerman 2012b). history stages of this cactus species indicate that seedlings The vegetation is classified as a subtropical dry forest, with a and juveniles are particularly susceptible to variations in local large proportion of species showing xeromorphic adaptations microclimatic conditions (Rojas-Sandoval and Meléndez- (Cintrón and Rogers 1991). Harrisia is one of three columnar Ackerman 2012a; 2012b). For this species, authors have cacti reported on the island (Rojas-Sandoval and Meléndez- shown that perennial native shrubs, such as Croton discolor Ackerman 2011a). Juveniles of Harrisia are unbranched, and Reynosia uncinata, act as nurse plants that provide suitable whereas adults usually are extensively branched and may microclimatic conditions that significantly improve survival, reach heights >2 m (Rojas-Sandoval and Meléndez-Ackerman establishment and growth of Harrisia seedlings (Rojas- 2011b). This species is an iteroparous night-flowering cactus Sandoval and Meléndez-Ackerman 2012a). While the com- that needs to set seed to propagate under natural conditions bined results suggest that abiotic factors are indeed important (Rojas-Sandoval and Meléndez-Ackerman 2009, 2011a). to the successful establishment and growth of Harrisia, we currently lack information on how
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